Body size and digestive system shape resource selection by ungulates: A cross-taxa test of the forage maturation hypothesis.

The forage maturation hypothesis (FMH) states that energy intake for ungulates is maximised when forage biomass is at intermediate levels. Nevertheless, metabolic allometry and different digestive systems suggest that resource selection should vary across ungulate species. By combining GPS relocations with remotely sensed data on forage characteristics and surface water, we quantified the effect of body size and digestive system in determining movements of 30 populations of hindgut fermenters (equids) and ruminants across biomes. Selection for intermediate forage biomass was negatively related to body size, regardless of digestive system. Selection for proximity to surface water was stronger for equids relative to ruminants, regardless of body size. To be more generalisable, we suggest that the FMH explicitly incorporate contingencies in body size and digestive system, with small-bodied ruminants selecting more strongly for potential energy intake, and hindgut fermenters selecting more strongly for surface water.

Predicting the continuum between corridors and barriers to animal movements using Step Selection Functions and Randomized Shortest Paths.

The loss, fragmentation and degradation of habitat everywhere on Earth prompts increasing attention to identifying landscape features that support animal movement (corridors) or impedes it (barriers). Most algorithms used to predict corridors assume that animals move through preferred habitat either optimally (e.g. least cost path) or as random walkers (e.g. current models), but neither extreme is realistic. We propose that corridors and barriers are two sides of the same coin and that animals experience landscapes as spatiotemporally dynamic corridor-barrier continua connecting (separating) functional areas where individuals fulfil specific ecological processes. Based on this conceptual framework, we propose a novel methodological approach that uses high-resolution individual-based movement data to predict corridor-barrier continua with increased realism. Our approach consists of two innovations. First, we use step selection functions (SSF) to predict friction maps quantifying corridor-barrier continua for tactical steps between consecutive locations. Secondly, we introduce to movement ecology the randomized shortest path algorithm (RSP) which operates on friction maps to predict the corridor-barrier continuum for strategic movements between functional areas. By modulating the parameter Ѳ, which controls the trade-off between exploration and optimal exploitation of the environment, RSP bridges the gap between algorithms assuming optimal movements (when Ѳ approaches infinity, RSP is equivalent to LCP) or random walk (when Ѳ → 0, RSP → current models). Using this approach, we identify migration corridors for GPS-monitored wild reindeer (Rangifer t. tarandus) in Norway. We demonstrate that reindeer movement is best predicted by an intermediate value of Ѳ, indicative of a movement trade-off between optimization and exploration. Model calibration allows identification of a corridor-barrier continuum that closely fits empirical data and demonstrates that RSP outperforms models that assume either optimality or random walk. The proposed approach models the multiscale cognitive maps by which animals likely navigate real landscapes and generalizes the most common algorithms for identifying corridors. Because suboptimal, but non-random, movement strategies are likely widespread, our approach has the potential to predict more realistic corridor-barrier continua for a wide range of species.

Harvest and decimation affect genetic drift and the effective population size in wild reindeer.

Harvesting and culling are methods used to monitor and manage wildlife diseases. An important consequence of these practices is a change in the genetic dynamics of affected populations that may threaten their long-term viability. The effective population size (N e) is a fundamental parameter for describing such changes as it determines the amount of genetic drift in a population. Here, we estimate N e of a harvested wild reindeer population in Norway. Then we use simulations to investigate the genetic consequences of management efforts for handling a recent spread of chronic wasting disease, including increased adult male harvest and population decimation. The N e/N ratio in this population was found to be 0.124 at the end of the study period, compared to 0.239 in the preceding 14 years period. The difference was caused by increased harvest rates with a high proportion of adult males (older than 2.5 years) being shot (15.2% in 2005-2018 and 44.8% in 2021). Increased harvest rates decreased N e in the simulations, but less sex biased harvest strategies had a lower negative impact. For harvest strategies that yield stable population dynamics, shifting the harvest from calves to adult males and females increased N e. Population decimation always resulted in decreased genetic variation in the population, with higher loss of heterozygosity and rare alleles with more severe decimation or longer periods of low population size. A very high proportion of males in the harvest had the most severe consequences for the loss of genetic variation. This study clearly shows how the effects of harvest strategies and changes in population size interact to determine the genetic drift of a managed population. The long-term genetic viability of wildlife populations subject to a disease will also depend on population impacts of the disease and how these interact with management actions.

New indicator of habitat functionality reveals high risk of underestimating trade-offs among sustainable development goals: The case of wild reindeer and hydropower.

Although biodiversity is crucial for Sustainable Development Goals (SDGs), following the current trajectory, we risk failing SDG 15. Using a new indicator quantifying the loss of functional habitat (habitat that is simultaneously suitable and well-connected), we show that the real impact of renewable energy is far larger than previously assumed. Specifically, we estimate that the construction of hydropower reservoirs in south Norway caused a loss of ca. 222 km2 of functional habitat for wild reindeer (Rangifer tarandus)-which is far larger than assumed based on land inundation indices (110 km2). Fully mitigating these impacts is challenging: scenario analyses reveal that the measures proposed by societal actors would yield only a fraction of the habitat lost (2-12 km2) and could cause trade-off risks with other SDGs. Using indices of functional connectivity is crucial for environmental impact assessments, as entire ecological networks for several species can be affected far beyond the reservoirs.

Habitat functionality: Integrating environmental and geographic space in niche modeling for conservation planning.

Niche modeling is typically used to assess the effects of anthropogenic land use and climate change on species distributions and to inform spatial conservation planning. These models focus on the suitability of local biotic and abiotic conditions for a species in environmental space (E-space). Although movements also affect species occurrence, efforts to formally integrate geographic space (G-space) into niche modeling have been hindered by the lack of comprehensive theoretical frameworks. We propose the "functional habitat" framework to define areas that are simultaneously of high quality in E-space, and functionally connected to other suitable habitats in G-space. Originating in metapopulation ecology, approaches have been developed to assess the amount of suitable connected habitats, based on the proximity between pairs of locations. Using network theory, which operates in topological space (T-space, defined by a network), we extended these metapopulation approaches to integrate movement constraints in G-space with niche modeling in E-space. We demonstrate the functional habitat framework using empirical data (GPS tracking and population monitoring) throughout the European wild mountain reindeer (Rangifer t. tarandus) distribution range. We show that functional habitat outperforms traditional suitability in explaining the species' distribution. This approach integrates effects from habitat loss and fragmentation for spatial conservation planning, and avoids overemphasizing small, inaccessible areas with locally suitable habitats. The functional habitat framework formally integrates biotic, abiotic, and movement constraints in niche modeling using network theory, thus opening a wide range of applications in spatial conservation planning.

Parasite Spillover from Domestic Sheep to Wild Reindeer-The Role of Salt Licks.

Attraction sites are important for environmental pathogen transmission and spillover. Yet, their role in wildlife disease dynamics is often poorly substantiated. Herein, we study the role of salt licks as potential attraction sites for the spillover of gastrointestinal parasites from domestic sheep to wild reindeer. Eggs from the introduced sheep nematode Nematodirus battus were found in faecal samples of both species, suggestive of spillover. DNA metabarcoding of soil, collected at salt licks, revealed that N. battus, in addition to Teladorsagia circumcincta, were the most frequently occurring parasitic nematodes, with a significantly higher prevalence of nematodal DNA in salt lick soil compared to soil from control sites nearby. The finding of similar DNA haplotypes of N. battus in sheep, reindeer, and salt lick soil supports the hypothesis of spillover to reindeer via salt licks. More detailed investigation of the genetic diversity of N. battus across these hosts is needed to draw firm conclusions. Infection with these sheep nematodes could potentially explain a recently observed decline in the calf recruitment rate of the Knutshø reindeer herd. This study also supports the hypothesized role of artificial salt licks as hot spots for the transmission of environmentally persistent pathogens and illustrates the importance of knowledge about such attraction points in the study of disease in free-roaming animals.

Optimal foraging by a large ungulate in an extreme environment: Wild mountain reindeer select snow-free feeding habitats in winter.

Optimal foraging models predict that individual animals will optimize net energy gain by intensifying forage activity and/or reducing forage energy cost. Then, the free distribution model predicts an animal's distribution in a patchy landscape will match the distribution of the resources. If not modified by other factors, such patterns may be expected to be particularly explicit in variable and extreme, forage-limited, and patchy environments, notably alpine and Arctic environments during winter. The large ungulate wild mountain reindeer (Rangifer tarandus tarandus) surviving in such environments is used as a model during the forage-limited winter season. The largest wild reindeer area in Western Europe (Hardangervidda, 8130 km2) is actively managed to sustain 10,000-12,000 wild reindeer. Since 2001, 104 different individuals have been GPS-tracked at 3-hr intervals. In winter, mountain reindeer may either choose to seek out and forage in patchy snow-free habitats, typically on top of wind-blown ridges, or use energy-demanding digging through the snow to reach ground forage (cratering). We use late April satellite data from Landsat 5 and 8 (30 × 30 m), airborne laser scanning subsampling (processed to 1 × 1 m grid), and topographic information (1 m resolution) derived from digital aerial photographs (0.25 × 0.25 m resolution) to delineate snow-free patches, constituting less than 694 km2. By overlaying recorded wild reindeer GPS positions winters 2001-2017 (188,942 positions), we document a strong positive selection for snow-free patches, which were used about four times more frequently than expected from a "random walk" model. On a daily basis, the preference for snow-free areas was slightly stronger in the evenings. In the sustainable management of wild mountain reindeer, the area of snow-free patches is an important predictor of winter forage availability and important winter source areas. It may be derived from remote sensing data.

High winter loads of Oestrid larvae and Elaphostrongylus rangiferi are associated with emaciation in wild reindeer calves.

The Oestrid flies Cephemyia trompe and Hypoderma tarandi and the nematode Elaphostrongylus rangiferi are important parasites of Rangifer spp. The larvae of Oestrid flies develop in the throat (C. trompe) and skin (H. tarandi) of their host during winter while E. rangiferi develop in the CNS. Oestrid pupation, and development of E. rangiferi larvae from first- (L1) to infective third- stage in the environment during summer are highly temperature dependent. We investigated the possible negative effects of these parasites on the winter body-condition of wild reindeer calves. Two year-classes (generations) of calf, born in a warm (2014) and cold (2015) summer respectively, were examined for changes in body condition between autumn and spring, in relation to the parasite load determined in the spring. The body condition in the autumn was assessed as carcass weight, while the body condition in the spring was assessed as carcass weight, supplemented by an evaluation of fat reserves in various bodily locations. Oestrids were counted directly whereas the E. rangiferi quantification was based on faecal counts of L1 larvae. The abundance of infections for Oestrids and E. rangiferi were significantly greater in the 2014 generation than in the 2015 generation. The mean carcass weight decreased between autumn and spring for the 2014 generation but increased in the 2015 generation. Emaciation in the spring was documented (fat reserve evaluation) in 42% and 7% of calves in the 2014 and 2015 cohorts, respectively. There was a significant correlation between high parasite load and the probability of emaciation. The mean summer temperature in 2014 was 2.6 °C higher than the mean for 2015, and 1.0 °C higher than the mean for the last 30-years. Our findings suggest that following a warm summer, high loads of Oestrids and E. rangiferi may cause emaciation and potentially deaths among the calves.

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.

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.

Elaphostrongylus and Dictyocaulus infections in Norwegian wild reindeer and red deer populations in relation to summer pasture altitude and climate.

Nematodes of the genera Elaphostrongylus and Dictyocaulus are associated with disease in semi-domesticated tundra reindeer and farmed red deer whereas less knowledge exists in the wild. Their first stage larvae (L1) develop to the infective third stage (L3) in the environment; Elaphostrongylus spp. within intermediate gastropod hosts and Dictyocaulus spp. as free-living larvae. Larval development of Elaphostrongylus is highly temperature dependent with a developmental minimum of 9-10 °C. Larval development of Dictyocaulus spp. may occur at low temperatures (5 °C) but the larvae are sensitive to desiccation. We examined the prevalence and intensity of Elaphostrongylus spp. and Dictyocaulus spp. infections in six wild reindeer and two wild red deer populations in relation to altitude, temperature and rainfall in their respective main summer pasture area over the 5 summers prior to sampling. The parasitological examination was based upon morphological identification of L1 in the faeces of hunted animals. Altitude was calculated from animal position data and temperature and precipitation by means of a nationwide gridded data set. Temperature decreased with increasing altitude, from 13.3 °C for the lowest located red deer population (300 m) to 6.1 °C for the highest located reindeer population (1400 m). No significant relationship between altitude and rainfall was identified. Elaphostrongylus spp. infection decreased in prevalence with increasing altitude, being identified in 89% of investigated samples from the lowest located population and in 3% of samples from the highest. The prevalence of Dictyocaulus spp. infection varied between 28 and 80% and no relationship with altitude was found. The intensity of Elaphostrongylus spp. infection was low in reindeer and moderate in red deer whereas the intensity of Dictyocaulus spp. infection was moderate in both species. Our results indicated that the climatic conditions in all areas studied were suitable for Dictyocaulus spp., whereas summer temperature was a restrictive factor for Elaphostrongylus sp. in reindeer.

Integrating data from multiple sources for insights into demographic processes: Simulation studies and proof of concept for hierarchical change-in-ratio models.

We developed a model for estimating demographic rates and population abundance based on multiple data sets revealing information about population age- and sex structure. Such models have previously been described in the literature as change-in-ratio models, but we extend the applicability of the models by i) using time series data allowing the full temporal dynamics to be modelled, by ii) casting the model in an explicit hierarchical modelling framework, and by iii) estimating parameters based on Bayesian inference. Based on sensitivity analyses we conclude that the approach developed here is able to obtain estimates of demographic rate with high precision whenever unbiased data of population structure are available. Our simulations revealed that this was true also when data on population abundance are not available or not included in the modelling framework. Nevertheless, when data on population structure are biased due to different observability of different age- and sex categories this will affect estimates of all demographic rates. Estimates of population size is particularly sensitive to such biases, whereas demographic rates can be relatively precisely estimated even with biased observation data as long as the bias is not severe. We then use the models to estimate demographic rates and population abundance for two Norwegian reindeer (Rangifer tarandus) populations where age-sex data were available for all harvested animals, and where population structure surveys were carried out in early summer (after calving) and late fall (after hunting season), and population size is counted in winter. We found that demographic rates were similar regardless whether we include population count data in the modelling, but that the estimated population size is affected by this decision. This suggest that monitoring programs that focus on population age- and sex structure will benefit from collecting additional data that allow estimation of observability for different age- and sex classes. In addition, our sensitivity analysis suggests that focusing monitoring towards changes in demographic rates might be more feasible than monitoring abundance in many situations where data on population age- and sex structure can be collected.

Physiologic evaluation of medetomidine-ketamine anesthesia in free-ranging Svalbard (Rangifer tarandus platyrhynchus) and wild Norwegian reindeer (Rangifer tarandus tarandus).

Previously published studies indicated that combinations of medetomidine and ketamine were effective for both Svalbard (Rangifer tarandus platyrhynchus) and wild Norwegian reindeer (Rangifer tarandus tarandus). Both previous studies indicated that reindeer were hypoxemic on the basis of pulse oximetry. We conducted a physiologic evaluation of these two protocols using arterial blood gases. Medetomidine (10 mg) and ketamine (200 mg) were administered by dart from the ground in Svalbard reindeer (October 2010) and from a helicopter for wild reindeer (March 2012). Of tested animals, all seven wild reindeer and five of seven Svalbard reindeer were hypoxemic before oxygen administration. Nasal oxygen insufflation (1 L/min for five Svalbard reindeer and one wild reindeer and 2 L/min for four wild reindeer) corrected hypoxemia in all cases evaluated. For reversal, all animals received 5 mg atipamezole per mg medetomidine intramuscularly.