Bear in mind! Bear presence and individual experience with calf survival shape the selection of calving sites in a long-lived solitary ungulate.

The careful selection of ungulate calving sites to improve offspring survival is vital in the face of predation. In general, there is limited knowledge to which degree predator presence and prey's individual experience shape the selection of calving sites. Predator presence influences the spatiotemporal risk of encountering a predator, while individual experiences with previous predation events shape perceived mortality risks. We used a multi-year movement dataset of a long-lived female ungulate (moose, Alces alces, n = 79) and associated calf survival to test how predator presence (i.e., encounter risk) and females' individual experiences with previous calf mortality events affected their calving site selection and site fidelity. Using data from areas with and without Scandinavian brown bear (Ursus arctos) predation, we compared females' calving site selection using individual-based analyses. Our findings suggest two things. First, bear presence influences calving site selection in this solitary living ungulate. Females in areas with bears were selected for higher shrub and tree cover and showed lower site fidelity than in the bear-free area. Second, the individual experience of calf loss changes females' selection the following year. Females with lost calves had a lower site fidelity compared to females with surviving calves. Our findings suggest that increased vegetation cover may be important for reducing encounter risk in bear areas, possibly by improving calf concealment. Lower site fidelity might represent a strategy to make the placement of calving sites less predictable for predators. We suggest that bear presence shapes both habitat selection and calving site fidelity in a long-lived animal, whereas the effect of individual experience with previous calf loss varies. We encourage further research on the relevance of female experience on the success of expressed anti-predator strategies during calving periods.

Sporadic cases of chronic wasting disease in old moose - an epidemiological study.

Transmissible spongiform encephalopathies or prion diseases comprise diseases with different levels of contagiousness under natural conditions. The hypothesis has been raised that the chronic wasting disease (CWD) cases detected in Nordic moose (Alces alces) may be less contagious, or not contagious between live animals under field conditions. This study aims to investigate the epidemiology of CWD cases detected in moose in Norway, Sweden and Finland using surveillance data from 2016 to 2022.In total, 18 CWD cases were detected in Nordic moose. All moose were positive for prion (PrPres) detection in the brain, but negative in lymph nodes, all were old (mean 16 years; range 12-20) and all except one, were female. Age appeared to be a strong risk factor, and the sex difference may be explained by few males reaching high age due to hunting targeting calves, yearlings and males.The cases were geographically scattered, distributed over 15 municipalities. However, three cases were detected in each of two areas, Selbu in Norway and Arjeplog-Arvidsjaur in Sweden. A Monte Carlo simulation approach was applied to investigate the likelihood of such clustering occurring by chance, given the assumption of a non-contagious disease. The empirical P-value for obtaining three cases in one Norwegian municipality was less than 0.05, indicating clustering. However, the moose in Selbu were affected by different CWD strains, and over a 6 year period with intensive surveillance, the apparent prevalence decreased, which would not be expected for an ongoing outbreak of CWD. Likewise, the three cases in Arjeplog-Arvidsjaur could also indicate clustering, but management practices promotes a larger proportion of old females and the detection of the first CWD case contributed to increased awareness and sampling.The results of our study show that the CWD cases detected so far in Nordic moose have a different epidemiology compared to CWD cases reported from North America and in Norwegian reindeer (Rangifer tarandus tarandus). The results support the hypothesis that these cases are less contagious or not contagious between live animals under field conditions. To enable differentiation from other types of CWD, we support the use of sporadic CWD (sCWD) among the names already in use.

Range-wide and temporal genomic analyses reveal the consequences of near-extinction in Swedish moose.

Ungulate species have experienced severe declines over the past centuries through overharvesting and habitat loss. Even if many game species have recovered thanks to strict hunting regulation, the genome-wide impacts of overharvesting are still unclear. Here, we examine the temporal and geographical differences in genome-wide diversity in moose (Alces alces) over its whole range in Sweden by sequencing 87 modern and historical genomes. We found limited impact of the 1900s near-extinction event but local variation in inbreeding and load in modern populations, as well as suggestion of a risk of future reduction in genetic diversity and gene flow. Furthermore, we found candidate genes for local adaptation, and rapid temporal allele frequency shifts involving coding genes since the 1980s, possibly due to selective harvesting. Our results highlight that genomic changes potentially impacting fitness can occur over short time scales and underline the need to track both deleterious and selectively advantageous genomic variation.

Estimating the neutron yield in a deuterium plasma with the JET neutron camera.

The JET neutron camera is a well-established detector system at JET, which has 19 sightlines each equipped with a liquid scintillator. The system measures a 2D profile of the neutron emission from the plasma. A first principle physics method is used to estimate the DD neutron yield that is based on JET neutron camera measurements and is independent of other neutron measurements. This paper details the data reduction techniques, models of the neutron camera, simulations of neutron transport, and detector responses used to this end. The estimate uses a simple parameterized model of the neutron emission profile. The method makes use of the JET neutron camera's upgraded data acquisition system. It also accounts for neutron scattering near the detectors and transmission through the collimator. These components together contribute to 9% of the detected neutron rate above a 0.5 MeVee energy threshold. Despite the simplicity of the neutron emission profile model, the DD neutron yield estimate falls on average within 10% agreement with a corresponding estimate from the JET fission chambers. The method can be improved by considering more advanced neutron emission profiles. It can also be expanded to estimate the DT neutron yield with the same methodology.

Increased summer temperature is associated with reduced calf mass of a circumpolar large mammal through direct thermoregulatory and indirect, food quality, pathways.

Climate change represents a growing ecological challenge. The (sub) arctic and boreal regions of the world experience the most rapid warming, presenting an excellent model system for studying how climate change affects mammals. Moose (Alces alces) are a particularly relevant model species with their circumpolar range. Population declines across the southern edge of this range are linked to rising temperatures. Using a long-term dataset (1988-1997, 2017-2019), we examine the relative strength of direct (thermoregulatory costs) and indirect (food quality) pathways linking temperature, precipitation, and the quality of two important food items (birch and fireweed) to variation in moose calf mass in northern Sweden. The direct effects of temperature consistently showed stronger relationships to moose calf mass than did the indirect effects. The proportion of growing season days where the temperature exceeded a 20 °C threshold showed stronger direct negative relationships to moose calf mass than did mean temperature values. Finally, while annual forb (fireweed) quality was more strongly influenced by temperature and precipitation than were perennial (birch) leaves, this did not translate into a stronger relationship to moose calf weight. The only indirect path with supporting evidence suggested that mean growing season temperatures were positively associated with neutral detergent fiber, which was, in turn, negatively associated with calf mass. While indirect impacts of climate change deserve further investigation, it is important to recognize the large direct impacts of temperature on cold-adapted species.

Body temperature patterns during pregnancy and parturition in moose.

Gestation and lactation have high energetic requirements. Up to three-fourths of the gestation period in moose (Alces alces) overlaps with the food-scarce period in winter. During this period, moose deal with the limited forage resources available through hypometabolism with decreased heart rate and body temperature (Tb). Body temperature is also an indicator of oestrus, pregnancy and parturition, which is well documented in several domestic species. In this study, we sought to determine if moose displayed a similar Tb pattern during pregnancy and parturition to domesticated ruminants, and if we could detect parturition by combining Tb and activity data. We studied the Tb pattern of 30 free-ranging adult female moose (≥1.5 years old), equipped with ruminal temperature loggers and GPS collars. We documented a 0.13-0.19°C higher Tb in pregnant compared to non-pregnant moose, depending on the study area with the Tb difference increasing along a south-north gradient, and a drop in Tb and in activity when parturition was imminent. Detection of parturition was highly successful when combining Tb and activity data with an accuracy of 91.5%. Our findings demonstrate that Tb responses to pregnancy and parturition in a wild capital-breeding ruminant are similar to those of domesticated ruminants.

Large mammal telomere length variation across ecoregions.

BACKGROUND: Telomere length provides a physiological proxy for accumulated stress in animals. While there is a growing consensus over how telomere dynamics and their patterns are linked to life history variation and individual experience, knowledge on the impact of exposure to different stressors at a large spatial scale on telomere length is still lacking. How exposure to different stressors at a regional scale interacts with individual differences in life history is also poorly understood. To better understand large-scale regional influences, we investigated telomere length variation in moose (Alces alces) distributed across three ecoregions. We analyzed 153 samples of 106 moose representing moose of both sexes and range of ages to measure relative telomere lengths (RTL) in white blood cells. RESULTS: We found that average RTL was significantly shorter in a northern (montane) and southern (sarmatic) ecoregion where moose experience chronic stress related to severe summer and winter temperatures as well as high anthropogenic land-use compared to the boreal region. Our study suggests that animals in the northern boreal forests, with relatively homogenous land use, are less disturbed by environmental and anthropogenic stressors. In contrast, animals in areas experiencing a higher rate of anthropogenic and environmental change experience increased stress. CONCLUSION: Although animals can often adapt to predictable stressors, our data suggest that some environmental conditions, even though predictable and ubiquitous, can generate population level differences of long-term stress. By measuring RTL in moose for the first time, we provide valuable insights towards our current understanding of telomere biology in free-ranging wildlife in human-modified ecosystems.

Integrating omics to characterize eco-physiological adaptations: How moose diet and metabolism differ across biogeographic zones.

With accelerated land conversion and global heating at northern latitudes, it becomes crucial to understand, how life histories of animals in extreme environments adapt to these changes. Animals may either adapt by adjusting foraging behavior or through physiological responses, including adjusting their energy metabolism or both. Until now, it has been difficult to study such adaptations in free-ranging animals due to methodological constraints that prevent extensive spatiotemporal coverage of ecological and physiological data.Through a novel approach of combining DNA-metabarcoding and nuclear magnetic resonance (NMR)-based metabolomics, we aim to elucidate the links between diets and metabolism in Scandinavian moose Alces alces over three biogeographic zones using a unique dataset of 265 marked individuals.Based on 17 diet items, we identified four different classes of diet types that match browse species availability in respective ecoregions in northern Sweden. Individuals in the boreal zone consumed predominantly pine and had the least diverse diets, while individuals with highest diet diversity occurred in the coastal areas. Males exhibited lower average diet diversity than females.We identified several molecular markers indicating metabolic constraints linked to diet constraints in terms of food availability during winter. While animals consuming pine had higher lipid, phospocholine, and glycerophosphocholine concentrations in their serum than other diet types, birch- and willow/aspen-rich diets exhibit elevated concentrations of several amino acids. The individuals with highest diet diversity had increased levels of ketone bodies, indicating extensive periods of starvation for these individuals.Our results show how the adaptive capacity of moose at the eco-physiological level varies over a large eco-geographic scale and how it responds to land use pressures. In light of extensive ongoing climate and land use changes, these findings pave the way for future scenario building for animal adaptive capacity.

Divergence in parturition timing and vegetation onset in a large herbivore-differences along a latitudinal gradient.

In northern environments, the period of access to high-quality forage is limited, exerting strong selective pressure to optimize the timing of parturition. We analysed timing and variation in moose (Alces alces) parturition dates of 555 females at 18 study sites across 12° of latitude (56-68° N, 1350 km) in Sweden. We found evidence for a spatial match of parturition timing to vegetation onset, but no evidence that moose adjust parturition to vegetation onset in a given year. We found a breakpoint at 64° N. Despite adaptation across latitudes, temporal divergences occurred. Females below 64° N calved after vegetation onset and females above 64° N calved before. Here, parturition before vegetation onset might be a strategy to optimize forage utilization time with the very short growing season. Highly seasonal environments such as at higher latitudes may make it advantageous to adapt parturition towards long-term climatic patterns by matching the most favourable period. Given the direction of temporal divergence, our study suggests that climate change may have less of an impact on moose parturition at northern latitudes than southern latitudes.

Varied diets, including broadleaved forage, are important for a large herbivore species inhabiting highly modified landscapes.

Diet quality is an important determinant of animal survival and reproduction, and can be described as the combination of different food items ingested, and their nutritional composition. For large herbivores, human landscape modifications to vegetation can limit such diet-mixing opportunities. Here we use southern Sweden's modified landscapes to assess winter diet mixtures (as an indicator of quality) and food availability as drivers of body mass (BM) variation in wild moose (Alces alces). We identify plant species found in the rumen of 323 moose harvested in Oct-Feb, and link variation in average calf BM among populations to diets and food availability. Our results show that variation in calf BM correlates with variation in diet composition, diversity, and food availability. A varied diet relatively rich in broadleaves was associated with higher calf BM than a less variable diet dominated by conifers. A diet high in shrubs and sugar/starch rich agricultural crops was associated with intermediate BM. The proportion of young production forest (0-15 yrs) in the landscape, an indicator of food availability, significantly accounted for variation in calf BM. Our findings emphasize the importance of not only diet composition and forage quantity, but also variability in the diets of large free-ranging herbivores.

Physiological and behavioural responses of moose to hunting with dogs.

Optimal management of hunted species requires an understanding of the impacts of hunting on both individual animal and population levels. Recent technological advancements in biologging enable us to obtain increasingly detailed information from free-ranging animals, covering longer periods of time, and providing the data needed to assess such impacts. In Sweden, more than 80 000 moose are harvested annually, mostly hunted with the use of baying dogs. The effects of this hunting method on animal welfare and stress are understudied. Here, we evaluated 6 real and 17 experimental hunting approaches with baying dogs [wearing global positioning system (GPS) collars] on 8 adult female moose equipped with ruminal temperature loggers, subcutaneous heart rate (HR) loggers and GPS collars with accelerometers. The obtained data were used to analyse the behavioural and physiological responses of moose to hunting with dogs. Successful experimental approaches (moose and dog were within 240 m for >10 min) resulted in higher maximum body temperature (Tb, 0.88°C higher) and a mean increase in HR of 24 bpm in moose at the day of the approach compared to the day after. The moose rested on average >90 min longer the day after the approach compared to the day of the approach. The moose travelled on average 4.2 km longer and had a 1.3 m/s higher maximum speed the day of the approach compared to the day after. Our results demonstrate that hunting with dogs increase moose energy expenditure and resting time (and consequently decrease time available for foraging) on an individual level. This could possibly affect body condition and reproduction rates if the hunting disturbances occur frequently.

Rate of Cooling in a Moose (Alces alces) Carcass.

Postmortem body temperature is used to estimate time of death in humans, but the available models are not validated for most nonhuman species. Here, we report that cooling in an adult female moose (Alces alces) equipped with a rumen temperature monitor was extremely slow, with a rumen temperature of 27-28 C as late as 40 h postmortem.

Effective thiafentanil immobilization and physiological responses of free-ranging moose (Alces alces) in northern Sweden.

OBJECTIVE: To evaluate clinical and physiological responses in moose to thiafentanil administration for immobilization. STUDY DESIGN: Cross-sectional clinical study. ANIMALS: Eleven (six males and five females) free-ranging adult moose (Alces alces). METHODS: Each moose was darted from a helicopter with 7.5 mg thiafentanil during March 2014 in northern Sweden. Physiological evaluation included vital signs and blood gases. Arterial blood was collected after induction and again after 10 minutes of intranasal oxygen administration and analyzed immediately with an i-STAT analyzer. A total of 10 mg naltrexone per milligram of thiafentanil was administered to all animals for reversal. Data were analyzed using descriptive statistics. RESULTS: All moose were sufficiently immobilized with a single dart injection. Induction occurred within 3 minutes in 10 of 11 moose. One individual became recumbent while crossing a river and naltrexone was immediately administered. Animals maintained sternal recumbency with their head raised and vital signs were stable. Nine of 10 moose were hypoxemic before oxygen administration, with seven becoming markedly hypoxemic [partial pressure of arterial oxygen (PaO2) between 40 and 59 mmHg (5.3-7.9 kPa)]. The PaO2 increased significantly between samples, but six moose remained hypoxemic despite therapy. Hypercapnia was seen in all moose, with eight having marked hypercapnia [partial pressure of arterial carbon dioxide (PaCO2) > 60 mmHg (>8.0 kPa)]. All moose were acidemic, with nine showing marked acidemia (pH < 7.20). The pH increased significantly with time and lactate decreased. Recoveries were rapid and uneventful, and all moose were living 6 months after capture. CONCLUSIONS: Thiafentanil provided rapid and sufficient immobilization of moose and its effects were rapidly reversed with naltrexone. As with other opioids, moose showed hypoxemia and varying degrees of respiratory and metabolic acidosis. Arterial oxygenation of moose improved following intranasal oxygen, but hypoxemia was not fully resolved despite therapy. CLINICAL RELEVANCE: Thiafentanil (7.5 mg per adult) is effective for immobilization of free-ranging moose. Supplemental oxygen may be of benefit when using this regimen; however, further investigation is required to confirm these results.

Discovery of SNPs for individual identification by reduced representation sequencing of moose (Alces alces).

Monitoring of wild animal populations is challenging, yet reliable information about population processes is important for both management and conservation efforts. Access to molecular markers, such as SNPs, enables population monitoring through genotyping of various DNA sources. We have developed 96 high quality SNP markers for individual identification of moose (Alces alces), an economically and ecologically important top-herbivore in boreal regions. Reduced representation libraries constructed from 34 moose were high-throughput de novo sequenced, generating nearly 50 million read pairs. About 50 000 stacks of aligned reads containing one or more SNPs were discovered with the Stacks pipeline. Several quality criteria were applied on the candidate SNPs to find markers informative on the individual level and well representative for the population. An empirical validation by genotyping of sequenced individuals and additional moose, resulted in the selection of a final panel of 86 high quality autosomal SNPs. Additionally, five sex-specific SNPs and five SNPs for sympatric species diagnostics are included in the panel. The genotyping error rate was 0.002 for the total panel and probability of identities were low enough to separate individuals with high confidence. Moreover, the autosomal SNPs were highly informative also for population level analyses. The potential applications of this SNP panel are thus many including investigations of population size, sex ratios, relatedness, reproductive success and population structure. Ideally, SNP-based studies could improve today's population monitoring and increase our knowledge about moose population dynamics.

A Unique Fatal Moose Attack Mimicking Homicide.

Fatalities caused by animal attacks are rare, but have the potential to mimic homicide. We present a case in which a moose attacked and killed a woman who was walking her dog in a forest. Autopsy showed widespread blunt trauma with a large laceration on one leg in which blades of grass were embedded. Flail chest was the cause of death. The case was initially conceived as homicide by means of a riding lawn mower. A review of the case by moose experts and analyses of biological trace material that proved to originate from moose, established the true source of injury. The dog probably provoked a moose, which, in response, stomped and gored the victim to death. The injuries resembled those previously reported from attacks by cattle and water buffalo. Fatal moose attacks constitute an extremely rare threat in boreal areas, but can be considered in traumatic deaths of unknown cause.

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.

Temporal patterns of moose-vehicle collisions with and without personal injuries.

Collisions with wild ungulates are an increasing traffic safety issue in boreal regions. Crashes involving smaller-bodied deer species usually lead to vehicle damage only, whereas collisions with a large animal, such as the moose, increase the risk of personal injuries. It is therefore important to understand both the factors affecting the number of moose-vehicle collisions (MVCs) and the underlying causes that turn an MVC into an accident involving personal injuries or fatalities. As a basis for temporal mitigation measures, we examined the annual and monthly variation of MVCs with and without personal injuries. Using a 22-year-long (1990-2011) time series from Finland, we tested the effect of moose population density and traffic volume on the yearly number of all MVCs and those leading to personal injuries. We also examined the monthly distribution of MVCs with and without personal injuries, and contrasted the Finnish findings with collision data from Sweden (years 2008-2010) and Norway (years 2008-2011). Both moose population abundance indices and traffic volume were positively related to the yearly variation in the number of MVCs in Finland. The proportion of MVCs involving personal injuries decreased during our 22-year study period. The monthly distribution of all MVCs peaked during the autumn or winter depending on country, while MVCs involving personal injury peaked in summer. Our study indicates that efforts to reduce MVCs involving personal injuries need to address driver awareness and attitudes during summer, despite most MVCs occurring in autumn or winter.

Quantifying Migration Behaviour Using Net Squared Displacement Approach: Clarifications and Caveats.

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.

Opportunities for the application of advanced remotely-sensed data in ecological studies of terrestrial animal movement.

Animal movement patterns in space and time are a central aspect of animal ecology. Remotely-sensed environmental indices can play a key role in understanding movement patterns by providing contiguous, relatively fine-scale data that link animal movements to their environment. Still, implementation of newly available remotely-sensed data is often delayed in studies of animal movement, calling for a better flow of information to researchers less familiar with remotely-sensed data applications. Here, we reviewed the application of remotely-sensed environmental indices to infer movement patterns of animals in terrestrial systems in studies published between 2002 and 2013. Next, we introduced newly available remotely-sensed products, and discussed their opportunities for animal movement studies. Studies of coarse-scale movement mostly relied on satellite data representing plant phenology or climate and weather. Studies of small-scale movement frequently used land cover data based on Landsat imagery or aerial photographs. Greater documentation of the type and resolution of remotely-sensed products in ecological movement studies would enhance their usefulness. Recent advancements in remote sensing technology improve assessments of temporal dynamics of landscapes and the three-dimensional structures of habitats, enabling near real-time environmental assessment. Online movement databases that now integrate remotely-sensed data facilitate access to remotely-sensed products for movement ecologists. We recommend that animal movement studies incorporate remotely-sensed products that provide time series of environmental response variables. This would facilitate wildlife management and conservation efforts, as well as the predictive ability of movement analyses. Closer collaboration between ecologists and remote sensing experts could considerably alleviate the implementation gap. Ecologists should not expect that indices derived from remotely-sensed data will be directly analogous to field-collected data and need to critically consider which remotely-sensed product is best suited for a given analysis.