COVID-19 related travel restrictions prevented numerous wildlife deaths on roads: A comparative analysis of results from 11 countries.

Millions of wild animals are killed annually on roads worldwide. During spring 2020, the volume of road traffic was reduced globally as a consequence of the COVID-19 pandemic. We gathered data on wildlife-vehicle collisions (WVC) from Czechia, Estonia, Finland, Hungary, Israel, Norway, Slovenia, Spain, Sweden, and for Scotland and England within the United Kingdom. In all studied countries WVC statistics tend to be dominated by large mammals (various deer species and wild boar), while information on smaller mammals as well as birds are less well recorded. The expected number of WVC for 2020 was predicted on the basis of 2015-2019 WVC time series representing expected WVC numbers under normal traffic conditions. Then, the forecasted and reported WVC data were compared. The results indicate varying levels of WVC decrease between countries during the COVID-19 related traffic flow reduction (CRTR). While no significant change was determined in Sweden, where the state-wide response to COVID-19 was the least intensive, a decrease as marked as 37.4% was identified in Estonia. The greatest WVC decrease, more than 40%, was determined during the first weeks of CRTR for Estonia, Spain, Israel, and Czechia. Measures taken during spring 2020 allowed the survival of large numbers of wild animals which would have been killed under normal traffic conditions. The significant effects of even just a few weeks of reduced traffic, help to highlight the negative impacts of roads on wildlife mortality and the need to boost global efforts of wildlife conservation, including systematic gathering of roadkill data.

Novel Type of Chronic Wasting Disease Detected in Moose (Alces alces), Norway.

Chronic wasting disease (CWD) persists in cervid populations of North America and in 2016 was detected for the first time in Europe in a wild reindeer in Norway. We report the detection of CWD in 3 moose (Alces alces) in Norway, identified through a large scale surveillance program. The cases occurred in 13-14-year-old female moose, and we detected an abnormal form of prion protein (PrPSc) in the brain but not in lymphoid tissues. Immunohistochemistry revealed that the moose shared the same neuropathologic phenotype, characterized by mostly intraneuronal deposition of PrPSc. This pattern differed from that observed in reindeer and has not been previously reported in CWD-infected cervids. Moreover, Western blot revealed a PrPSc type distinguishable from previous CWD cases and from known ruminant prion diseases in Europe, with the possible exception of sheep CH1641. These findings suggest that these cases in moose represent a novel type of CWD.

Movement is the glue connecting home ranges and habitat selection.

Animal space use has been studied by focusing either on geographic (e.g. home ranges, species' distribution) or on environmental (e.g. habitat use and selection) space. However, all patterns of space use emerge from individual movements, which are the primary means by which animals change their environment. Individuals increase their use of a given area by adjusting two key movement components: the duration of their visit and/or the frequency of revisits. Thus, in spatially heterogeneous environments, animals exploit known, high-quality resource areas by increasing their residence time (RT) in and/or decreasing their time to return (TtoR) to these areas. We expected that spatial variation in these two movement properties should lead to observed patterns of space use in both geographic and environmental spaces. We derived a set of nine predictions linking spatial distribution of movement properties to emerging space-use patterns. We predicted that, at a given scale, high variation in RT and TtoR among habitats leads to strong habitat selection and that long RT and short TtoR result in a small home range size. We tested these predictions using moose (Alces alces) GPS tracking data. We first modelled the relationship between landscape characteristics and movement properties. Then, we investigated how the spatial distribution of predicted movement properties (i.e. spatial autocorrelation, mean, and variance of RT and TtoR) influences home range size and hierarchical habitat selection. In landscapes with high spatial autocorrelation of RT and TtoR, a high variation in both RT and TtoR occurred in home ranges. As expected, home range location was highly selective in such landscapes (i.e. second-order habitat selection); RT was higher and TtoR lower within the selected home range than outside, and moose home ranges were small. Within home ranges, a higher variation in both RT and TtoR was associated with higher selectivity among habitat types (i.e. third-order habitat selection). Our findings show how patterns of geographic and environmental space use correspond to the two sides of a coin, linked by movement responses of individuals to environmental heterogeneity. By demonstrating the potential to assess the consequences of altering RT or TtoR (e.g. through human disturbance or climatic changes) on home range size and habitat selection, our work sets the basis for new theoretical and methodological advances in movement ecology.

Understanding scales of movement: animals ride waves and ripples of environmental change.

Animal movements are the primary behavioural adaptation to spatiotemporal heterogeneity in resource availability. Depending on their spatiotemporal scale, movements have been categorized into distinct functional groups (e.g. foraging movements, dispersal, migration), and have been studied using different methodologies. We suggest striving towards the development of a coherent framework based on the ultimate function of all movement types, which is to increase individual fitness through an optimal exploitation of resources varying in space and time. We developed a novel approach to simultaneously study movements at different spatiotemporal scales based on the following proposed theory: the length and frequency of animal movements are determined by the interaction between temporal autocorrelation in resource availability and spatial autocorrelation in changes in resource availability. We hypothesized that for each time interval the spatiotemporal scales of moose Alces alces movements correspond to the spatiotemporal scales of variation in the gains derived from resource exploitation when taking into account the costs of movements (represented by their proxies, forage availability NDVI and snow depth respectively). The scales of change in NDVI and snow were quantified using wave theory, and were related to the scale of moose movement using linear mixed models. In support of the proposed theory we found that frequent, smaller scale movements were triggered by fast, small-scale ripples of changes, whereas infrequent, larger scale movements matched slow, large-scale waves of change in resource availability. Similarly, moose inhabiting ranges characterized by larger scale waves of change in the onset of spring migrated longer distances. We showed that the scales of movements are driven by the scales of changes in the net profitability of trophic resources. Our approach can be extended to include drivers of movements other than trophic resources (e.g. population density, density of related individuals, predation risk) and may facilitate the assessment of the impact of environmental changes on community dynamics and conservation.

Coprophagy in moose: A first observation.

Coprophagy, the eating of feces, has been documented in a wide range of species but appears to be rare or difficult to detect in deer (Cervidae). Here, we report the first observation of coprophagy in moose Alces alces, which was recorded using camera collars on free-ranging moose in Norway. The footage shows an instance of allocoprophagy by an adult female moose in spring (May). We summarize the current knowledge about coprophagy in deer and briefly discuss potential drivers and possible implications for disease transmission. Further research is needed to determine whether coprophagy occurs frequently in moose and whether this behavior is positive (e.g., increased intake of nutrients) or negative (increased infection by parasites or pathogens).

Faecal metabarcoding provides improved detection and taxonomic resolution for non-invasive monitoring of gastrointestinal nematode parasites in wild moose populations.

BACKGROUND: Although wild ungulate populations are heavily monitored throughout Europe, we understand little of how parasites affect population dynamics, and there is no systematic, long-term monitoring of parasite diversity and parasite loads. Such monitoring is in part hampered by a lack of time- and cost-effective assay methodologies with high sensitivity and good taxonomic resolution. DNA metabarcoding has been successfully used to characterize the parasitic nemabiome with high taxonomic resolution in a variety of wild and domestic hosts. However, in order to implement this technique in large-scale, potentially non-invasive monitoring of gastrointestinal parasitic nematodes (GIN), protocol optimization is required to maximize biodiversity detection, whilst maintaining time- and cost-effectiveness. METHODS: Faecal samples were collected from a wild moose population and GIN communities were characterized and quantified using both parasitological techniques (egg and larva counting) and DNA metabarcoding of the ITS2 region of rDNA. Three different isolation methods were compared that differed in the volume of starting material and cell lysis method. RESULTS: Similar nematode faunas were recovered from all samples using both parasitological and metabarcoding methods, and the approaches were largely congruent. However, metabarcoding assays showed better taxonomic resolution and slightly higher sensitivity than egg and larvae counts. The metabarcoding was not strictly quantitative, but the proportion of target nematode sequences recovered was correlated with the parasitologically determined parasite load. Species detection rates in the metabarcoding assays were maximized using a DNA isolation method that included mechanical cell disruption and maximized the starting material volume. CONCLUSIONS: DNA metabarcoding is a promising technique for the non-invasive, large-scale monitoring of parasitic GINs in wild ungulate populations, owing to its high taxonomic resolution, increased assay sensitivity, and time- and cost-effectiveness. Although metabarcoding is not a strictly quantitative method, it may nonetheless be possible to create a management- and conservation-relevant index for the host parasite load from this data. To optimize the detection rates and time- and cost-effectiveness of metabarcoding assays, we recommend choosing a DNA isolation method that involves mechanical cell disruption and maximizes the starting material volume.

Moose in our neighborhood: Does perceived hunting risk have cascading effects on tree performance in vicinity of roads and houses?

Like large carnivores, hunters both kill and scare ungulates, and thus might indirectly affect plant performance through trophic cascades. In this study, we hypothesized that intensive hunting and enduring fear of humans have caused moose and other forest ungulates to partly avoid areas near human infrastructure (perceived hunting risk), with positive cascading effects on recruitment of trees. Using data from the Norwegian forest inventory, we found decreasing browsing pressure and increasing tree recruitment in areas close to roads and houses, where ungulates are more likely to encounter humans. However, although browsing and recruitment were negatively related, reduced browsing was only responsible for a small proportion of the higher tree recruitment near human infrastructure. We suggest that the apparently weak cascading effect occurs because the recorded browsing pressure only partly reflects the long-term browsing intensity close to humans. Accordingly, tree recruitment was also related to the density of small trees 5-10 years earlier, which was higher close to human infrastructure. Hence, if small tree density is a product of the browsing pressure in the past, the cascading effect is probably stronger than our estimates suggest. Reduced browsing near roads and houses is most in line with risk avoidance driven by fear of humans (behaviorally mediated), and not because of excessive hunting and local reduction in ungulate density (density mediated).

A model-driven approach to quantify migration patterns: individual, regional and yearly differences.

1. Animal migration has long intrigued scientists and wildlife managers alike, yet migratory species face increasing challenges because of habitat fragmentation, climate change and over-exploitation. Central to the understanding migratory species is the objective discrimination between migratory and nonmigratory individuals in a given population, quantifying the timing, duration and distance of migration and the ability to predict migratory movements. 2. Here, we propose a uniform statistical framework to (i) separate migration from other movement behaviours, (ii) quantify migration parameters without the need for arbitrary cut-off criteria and (iii) test predictability across individuals, time and space. 3. We first validated our novel approach by simulating data based on established theoretical movement patterns. We then formulated the expected shapes of squared displacement patterns as nonlinear models for a suite of movement behaviours to test the ability of our method to distinguish between migratory movement and other movement types. 4. We then tested our approached empirically using 108 wild Global Positioning System (GPS)-collared moose Alces alces in Scandinavia as a study system because they exhibit a wide range of movement behaviours, including resident, migrating and dispersing individuals, within the same population. Applying our approach showed that 87% and 67% of our Swedish and Norwegian subpopulations, respectively, can be classified as migratory. 5. Using nonlinear mixed effects models for all migratory individuals we showed that the distance, timing and duration of migration differed between the sexes and between years, with additional individual differences accounting for a large part of the variation in the distance of migration but not in the timing or duration. Overall, the model explained most of the variation (92%) and also had high predictive power for the same individuals over time (69%) as well as between study populations (74%). 6. The high predictive ability of the approach suggests that it can help increase our understanding of the drivers of migration and could provide key quantitative information for understanding and managing a broad range of migratory species.

Harvest strategies for the elimination of low prevalence wildlife diseases.

The intensive harvesting of hosts is often the only practicable strategy for controlling emerging wildlife diseases. Several harvesting approaches have been explored theoretically with the objective of lowering transmission rates, decreasing the transmission period or specifically targeting spatial disease clusters or high-risk demographic groups. Here, we present a novel model-based approach to evaluate alternative harvest regimes, in terms of demographic composition and rates, intended to increase the probability to remove all infected individuals in the population during the early phase of an outbreak. We tested the utility of the method for the elimination of chronic wasting disease based on empirical data for reindeer (Rangifer tarandus) in Norway, in populations with (Nordfjella) and without (Hardangervidda) knowledge about exact disease prevalence and population abundance. Low and medium harvest intensities were unsuccessful in eliminating the disease, even at low prevalence. High-intensity harvesting had a high likelihood of eliminating the disease, but probability was strongly influenced by the disease prevalence. We suggest that the uncertainty about disease prevalence can be mitigated by using an adaptive management approach: forecast from models after each harvest season with updated data, derive prevalence estimates and forecast further harvesting. We identified the problems arising from disease surveillance with large fluctuations in harvesting pressure and hence sample sizes. The elimination method may be suitable for pathogens that cause long-lasting infections and with slow epidemic growth, but the method should only be attempted if there is a low risk of reinfection, either by a new disease introduction event (e.g. dispersing hosts) or due to environmental reservoirs. Our simulations highlighted the short time window when such a strategy is likely to be successful before approaching near complete eradication of the population.

Identifying and correcting spatial bias in opportunistic citizen science data for wild ungulates in Norway.

Many publications make use of opportunistic data, such as citizen science observation data, to infer large-scale properties of species' distributions. However, the few publications that use opportunistic citizen science data to study animal ecology at a habitat level do so without accounting for spatial biases in opportunistic records or using methods that are difficult to generalize. In this study, we explore the biases that exist in opportunistic observations and suggest an approach to correct for them. We first examined the extent of the biases in opportunistic citizen science observations of three wild ungulate species in Norway by comparing them to data from GPS telemetry. We then quantified the extent of the biases by specifying a model of the biases. From the bias model, we sampled available locations within the species' home range. Along with opportunistic observations, we used the corrected availability locations to estimate a resource selection function (RSF). We tested this method with simulations and empirical datasets for the three species. We compared the results of our correction method to RSFs obtained using opportunistic observations without correction and to RSFs using GPS-telemetry data. Finally, we compared habitat suitability maps obtained using each of these models. Opportunistic observations are more affected by human access and visibility than locations derived from GPS telemetry. This has consequences for drawing inferences about species' ecology. Models naïvely using opportunistic observations in habitat-use studies can result in spurious inferences. However, sampling availability locations based on the spatial biases in opportunistic data improves the estimation of the species' RSFs and predicted habitat suitability maps in some cases. This study highlights the challenges and opportunities of using opportunistic observations in habitat-use studies. While our method is not foolproof it is a first step toward unlocking the potential of opportunistic citizen science data for habitat-use studies.

Distribution, prevalence and intensity of moose nose bot fly (Cephenemyia ulrichii) larvae in moose (Alces alces) from Norway.

High host density combined with climate change may lead to invasion of harmful parasites in cervid (host) populations. Bot flies (Diptera: Oestridae) are a group of ectoparasites that may have strong impact on their hosts, but data on the current distribution, prevalence and intensity of the moose nose bot fly (Cephenemyia ulrichii) in Scandinavia are lacking. We estimated prevalence and intensity of nose bot fly larvae in 30 moose from southern and 79 moose from central Norway. All larvae detected were identified as the moose nose bot fly. We found surprisingly high prevalence in these areas, which are up to 1300 km south-southwest of the first published location in Norway and west of the distribution in Sweden. Prevalence (0.44-1.00) was higher in areas with higher moose density. Parasite intensity in hunter killed moose was higher in central Norway (mean 5.7) than southern Norway (mean 2.9), and in both regions higher in calves and yearlings than adults. Fallen moose had higher parasite intensity (mean 9.8) compared to hunter killed moose in the subsample from central Norway, suggesting a link to host condition or behavior. Our study provides evidence of parasite range expansion, and establishing monitoring appears urgent to better understand impact on host populations.

The unique spatial ecology of human hunters.

Human hunters are described as 'superpredators' with a unique ecology. Chronic wasting disease among cervids and African swine fever among wild boar are emerging wildlife diseases in Europe, with huge economic and cultural repercussions. Understanding hunter movements at broad scales has implications for how to control the spread of these diseases. Here we show, based on analysis of the settlement patterns and movements of hunters of reindeer (n = 9,685), red deer (n = 47,845), moose (n = 60,365) and roe deer (n = 42,530) from across Norway (2001-2017), that hunter density was more closely linked to human density than prey density and that hunters were largely migratory, aggregated with increasing regional prey densities and often used dogs. Hunter movements extended across Europe and to other continents. Our results provide extensive evidence that the broad-scale movements and residency patterns of postindustrial hunters relative to their prey differ from those of large carnivores.

Antler cannibalism in reindeer.

Prion diseases constitute a class of invariably fatal and degenerative encephalopathies. Chronic Wasting Disease (CWD) is a contagious prion disease among cervids, which is spreading and causing marked population declines in USA and Canada. The first outbreak of CWD in Europe was discovered in a reindeer population in Norway in 2016. In the worst-case scenario with continental-wide spreading of CWD in Eurasia, an annual harvest of around 4 million cervids is at stake only in Europe, with huge economic and cultural significance. An in situ origin of CWD was suspected, and it appear urgent to identify the likely cause to prevent future emergences. Here, we document the novel phenomenon of extensive antler cannibalism prior to shedding among reindeer in the CWD-infected population. The extent of antler cannibalism increased over the last decades when CWD emerged, and included ingestion of vascularized antlers. Ingestion of tissues from conspecifics is a risk factor for the emergence of prion diseases, where the presence of extensive antler cannibalism opens the intriguing possibility of a 'Kuru-analogue' origin of CWD among the reindeer in Europe. Based on general insight on pathology of prion diseases and strain selection processes, we propose an hypothesis for how contagious CWD may emerge from sporadic CWD under the unique epidemiological conditions we document here. More research is required to document the presence of prions in reindeer antlers, and whether antler cannibalism actually led to a strain selection process and the emergence of a contagious form of CWD from a sporadic form of CWD.

Hunting strategies to increase detection of chronic wasting disease in cervids.

The successful mitigation of emerging wildlife diseases may involve controversial host culling. For livestock, 'preemptive host culling' is an accepted practice involving the removal of herds with known contact to infected populations. When applied to wildlife, this proactive approach comes in conflict with biodiversity conservation goals. Here, we present an alternative approach of 'proactive hunting surveillance' with the aim of early disease detection that simultaneously avoids undesirable population decline by targeting demographic groups with (1) a higher likelihood of being infected and (2) a lower reproductive value. We applied this harvesting principle to populations of reindeer to substantiate freedom of chronic wasting disease (CWD) infection. Proactive hunting surveillance reached 99% probability of freedom from infection (<4 reindeer infected) within 3-5 years, in comparison to ~10 years using ordinary harvest surveillance. However, implementation uncertainties linked to social issues appear challenging also with this kind of host culling.

Chronic wasting disease associated with prion protein gene (PRNP) variation in Norwegian wild reindeer (Rangifer tarandus).

The emergence of CWD in Europe in 2016 and the first natural infection in wild reindeer warranted disease management. This led to the testing of 2424 hunted or culled reindeer during 2016-2018, from the infected subpopulation in the Nordfjella mountain range in Southern Norway. To identify any association between PRNP variation and CWD susceptibility, we characterized the open reading frame of the PRNP gene in 19 CWD positive reindeer and in 101 age category- and sex-matched CWD negative controls. Seven variant positions were identified: 6 single nucleotide variants (SNVs) and a 24 base pair (bp) deletion located between nucleotide position 238 and 272, encoding four instead of five octapeptide repeats. With a single exception, all variant positions but one were predicted to be non-synonymous. The synonymous SNV and the deletion are novel in reindeer. Various combinations of the non-synonymous variant positions resulted in the identification of five PRNP alleles (A-E) that structured into 14 genotypes. We identified an increased CWD risk in reindeer carrying two copies of the most common allele, A, coding for serine in position 225 (Ser225) and in those carrying allele A together with the 24 bp deletion.

Plasma lactate concentrations in free-ranging moose (Alces alces) immobilized with etorphine.

OBJECTIVE: To investigate plasma lactate concentrations of etorphine-immobilized moose in relation to environmental, temporal and physiological parameters. STUDY DESIGN: Prospective clinical study. ANIMALS: Fourteen female and five male moose (Alces alces), estimated age range 1-7 years. METHODS: The moose were darted from a helicopter with 7.5 mg etorphine per animal using projectile syringes and a dart gun. Once immobilized, the moose were approached, a venous blood sample was obtained and vital signs including pulse oximetry were recorded. Diprenorphine was administered to reverse the effects of etorphine. Timing of events, ambient temperature and snow depth were recorded. Blood samples were cooled and centrifuged before plasma was harvested and frozen. The plasma was thawed later and lactate analysed. Data were analysed using descriptive statistics and regression analysis. RESULTS: All animals recovered uneventfully and were alive 12 weeks after immobilization. Mean +/- SD plasma lactate was found to be 9.2 +/- 2.1 mmol L(-1). Plasma lactate concentrations were related positively to snow depth and negatively to time from induction of immobilization to blood sampling. The model that best described the variability in plasma lactate concentrations used induction time (time from firing the dart to the moose being immobilized). The second best model included induction time and snow depth. CONCLUSIONS AND CLINICAL RELEVANCE: Plasma lactate concentrations in these etorphine-immobilized moose were in the range reported for other immobilized wild ruminants. Decreasing induction time, which may be related to a more profound etorphine effect, and increasing snow depth possibly may increase plasma lactate concentrations in etorphine-immobilized moose.

Infection prevalence and ecotypes of Anaplasma phagocytophilum in moose Alces alces, red deer Cervus elaphus, roe deer Capreolus capreolus and Ixodes ricinus ticks from Norway.

BACKGROUND: The geographical expansion of the tick Ixodes ricinus in northern Europe is a serious concern for animal and human health. The pathogen Anaplasma phagocytophilum is transmitted by ticks and causes emergences of tick-borne fever (anaplasmosis) in livestock. The transmission dynamics of the different ecotypes of A. phagocytophilum in the ecosystems is only partly determined. Red deer and roe deer contribute to circulation of different ecotypes of A. phagocytophilum in continental Europe, while the role of moose for circulation of different ecotypes is not fully established but an important issue in northern Europe. METHODS: We determined infection prevalence and ecotypes of A. phagocytophilum in moose (n = 111), red deer (n = 141), roe deer (n = 28) and questing ticks (n = 9241) in Norway. RESULTS: As previously described, red deer was exclusively linked to circulation of ecotype I, while roe deer was exclusively linked to circulation of ecotype II. Surprisingly, we found 58% ecotype I (n = 19) and 42% of ecotype II (n = 14) in moose. Both ecotypes were found in questing ticks in areas with multiple cervid species present, while only ecotype I was found in ticks in a region with only red deer present. Hence, the geographical distribution of ecotypes in ticks followed the distribution of cervid species present in a given region and their link to ecotype I and II. CONCLUSIONS: Moose probably function as reservoirs for both ecotype I and II, indicating that the ecotypes of A. phagocytophilum are not entirely host-specific and have overlapping niches. The disease hazard depends also on both host abundance and the number of immature ticks fed by each host. Our study provides novel insights in the northern distribution and expansion of tick-borne fever.

First Detection of Chronic Wasting Disease in a Wild Red Deer (Cervus elaphus) in Europe.

Chronic wasting disease (CWD) is a fatal contagious prion disease naturally occurring in cervids in North America. In 2016, CWD was detected in wild reindeer (Rangifer tarandus) and moose (Alces alces) in Norway. Here, we report the first known naturally infected wild Norwegian red deer (Cervus elaphus).

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.