BACKGROUND: Movement links the distribution of habitats with the social environment of animals using those habitats. Despite the links between movement, habitat selection, and socioecology, their integration remains a challenge due to lack of shared vocabulary across fields, methodological gaps, and the implicit (rather than explicit) historical development of theory in the fields of social and spatial ecology. Given these challenges can be addressed, opportunity for further study will provide insight about the links between social, spatial, and movement ecology. Here, our objective was to disentangle the roles of habitat selection and social association as drivers of movement in caribou (Rangifer tarandus).
METHODS: To accomplish our objective, we modelled the relationship between collective movement and selection of foraging habitats using socially informed integrated step selection function (iSSF). Using iSSF, we modelled the effect of social processes, i.e., nearest neighbour distance and social preference, and movement behaviour on patterns of habitat selection.
RESULTS: By unifying social network analysis with iSSF, we identified movement-dependent social association, where individuals took shorter steps in lichen habitat and foraged in close proximity to more familiar individuals.
CONCLUSIONS: Our study demonstrates that social preference is context-dependent based on habitat selection and foraging behaviour. We therefore surmise that habitat selection and social association are drivers of collective movement, such that movement is the glue between habitat selection and social association. Here, we put these concepts into practice to demonstrate that movement is the glue connecting individual habitat selection to the social environment.

Abundance estimation is frequently an objective of conservation and monitoring initiatives for threatened and other managed populations. While abundance estimation via capture-mark-recapture or spatially explicit capture-recapture is now common, such approaches are logistically challenging and expensive for species such as boreal caribou (Rangifer tarandus), which inhabit remote regions, are widely dispersed, and exist at low densities. Fortunately, the recently developed \'close-kin mark-recapture\' (CKMR) framework, which uses the number of kin pairs obtained within a sample to generate an abundance estimate, eliminates the need for multiple sampling events. As a result, some caribou managers are interested in using this method to generate an abundance estimate from a single, non-invasive sampling event for caribou populations. We conducted a simulation study using realistic boreal caribou demographic rates and population sizes to assess how population size and the proportion of the population surveyed impact the accuracy and precision of single-survey CKMR-based abundance estimates. Our results indicated that abundance estimates were biased and highly imprecise when very small proportions of the population were sampled, regardless of the population size. However, the larger the population size, the smaller the required proportion of the population surveyed to generate both accurate and reasonably precise estimates. Additionally, we also present a case study in which we used the CKMR framework to generate annual female abundance estimates for a small caribou population in Jasper National Park, Alberta, Canada, from 2006 to 2015 and compared them to existing published capture-mark-recapture-based estimates. Both the accuracy and precision of the annual CKMR-based abundance estimates varied across years and were sensitive to the proportion of pairwise kinship comparisons which yielded a mother-offspring pair. Taken together, our study demonstrates that it is possible to generate CKMR-based abundance estimates from a single sampling event for small caribou populations, so long as a sufficient sampling intensity can be achieved.

Reindeer, called caribou in North America, has a circumpolar distribution and all extant populations belong to the same species (Rangifer tarandus). It has survived the Holocene thanks to its immense adaptability and successful coexistence with humans in different forms of hunting and herding cultures. Here, we examine the paternal and maternal history of Rangifer based on robust Y-chromosomal and mitochondrial DNA (mtDNA) trees representing Eurasian tundra reindeer, Finnish forest reindeer, Svalbard reindeer, Alaska tundra caribou, and woodland caribou. We first assembled Y-chromosomal contigs, representing 1.3 Mb of single-copy Y regions. Based on 545 Y-chromosomal and 458 mtDNA SNPs defined in 55 males, maximum parsimony trees were created. We observed two well separated clades in both phylogenies: the \"EuroBeringian clade\" formed by animals from Arctic Islands, Eurasia, and a few from North America and the \"North American clade\" formed only by caribou from North America. The time calibrated Y tree revealed an expansion and dispersal of lineages across continents after the Last Glacial Maximum. We show for the first time unique paternal lineages in Svalbard reindeer and Finnish forest reindeer and reveal a circumscribed Y haplogroup in Fennoscandian tundra reindeer. The Y chromosome in domesticated reindeer is markedly diverse indicating that several male lineages have undergone domestication and less intensive selection on males. This study places R. tarandus onto the list of species with resolved Y and mtDNA phylogenies and builds the basis for studies of the distribution and origin of paternal and maternal lineages in the future.

Competition for resources and space can drive forage selection of large herbivores from the bite through the landscape scale. Animal behaviour and foraging patterns are also influenced by abiotic and biotic factors. Fine-scale mechanisms of density-dependent foraging at the bite scale are likely consistent with density-dependent behavioural patterns observed at broader scales, but few studies have directly tested this assertion. Here, we tested if space use intensity, a proxy of spatiotemporal density, affects foraging mechanisms at fine spatial scales similarly to density-dependent effects observed at broader scales in caribou. We specifically assessed how behavioural choices are affected by space use intensity and environmental processes using behavioural state and forage selection data from caribou (Rangifer tarandus granti) observed from GPS video-camera collars using a multivariate discrete-choice modelling framework. We found that the probability of eating shrubs increased with increasing caribou space use intensity and cover of Salix spp. shrubs, whereas the probability of eating lichen decreased. Insects also affected fine-scale foraging behaviour by reducing the overall probability of eating. Strong eastward winds mitigated negative effects of insects and resulted in higher probabilities of eating lichen. At last, caribou exhibited foraging functional responses wherein their probability of selecting each food type increased as the availability (% cover) of that food increased. Space use intensity signals of fine-scale foraging were consistent with density-dependent responses observed at larger scales and with recent evidence suggesting declining reproductive rates in the same caribou population. Our results highlight potential risks of overgrazing on sensitive forage species such as lichen. Remote investigation of the functional responses of foraging behaviours provides exciting future applications where spatial models can identify high-quality habitats for conservation.

Brucellosis is an important zoonotic disease affecting animals and subsistence harvesters in the circumarctic. We investigated recent trends (2015-2022) of brucellosis seropositivity in caribou (Rangifer tarandus) and muskoxen (Ovibos moschatus) in the Central Canadian Arctic by using data from community-based wildlife health surveillance programs. The overall sample prevalence of Brucella antibodies was 10.0% (n = 271) in muskoxen and 15.5% (n = 277) in caribou. Sample seroprevalence in muskoxen varied geographically with an increasing trend of exposure on NW Victoria Island (from 0% to 36.8% between 2016 and 2022; Kendall tau = 0.283, p = 0.001). The presence of Brucella suis biovar 4 was confirmed by culture from clinical cases in this area. Our results indicate that Brucella suis biovar 4 continues to circulate in the Central Canadian Arctic in caribou and muskoxen and may be now circulating in muskoxen independently from caribou. These findings highlight the need to better understand the ecology and drivers of brucellosis emergence in Arctic multi-host systems.

Reindeer in the Arctic seasonally suppress daily circadian patterns of behavior present in most animals.1 In humans and mice, even when all daily behavioral and environmental influences are artificially suppressed, robust endogenous rhythms of metabolism governed by the circadian clock persist and are essential to health.2,3 Disrupted rhythms foster metabolic disorders and weight gain.4 To understand circadian metabolic organization in reindeer, we performed behavioral measurements and untargeted metabolomics from blood plasma samples taken from Eurasian tundra reindeer (Rangifer tarandus tarandus) across 24 h at 2-h intervals in four seasons. Our study confirmed the absence of circadian rhythms of behavior under constant darkness in the Arctic winter and constant daylight in the Arctic summer, as reported by others.1 We detected and measured the intensity of 893 metabolic features in all plasma samples using untargeted ultra-high-performance liquid chromatography-mass spectrometry (UPLC-MS). A core group of metabolites (66/893 metabolic features) consistently displayed 24-h rhythmicity. Most metabolites displayed a robust 24-h rhythm in winter and spring but were arrhythmic in summer and fall. Half of all measured metabolites displayed ultradian sleep-wake dependence in summer. Irrespective of the arrhythmic behavior, metabolism is rhythmic (24 h) in seasons of low food availability, potentially favoring energy efficiency. In seasons of food abundance, 24-h rhythmicity in metabolism is drastically reduced, again irrespective of behavioral rhythms, potentially fostering weight gain.

High intra-specific genetic diversity is associated with adaptive potential, which is key for resilience to global change. However, high variation may also support deleterious alleles through genetic load, thereby increasing the risk of inbreeding depression if population sizes decrease. Purging of deleterious variation has been demonstrated in some threatened species. However, less is known about the costs of declines and inbreeding in species with large population sizes and high genetic diversity even though this encompasses many species globally that are expected to undergo population declines. Caribou is a species of ecological and cultural significance in North America with a wide distribution supporting extensive phenotypic variation but with some populations undergoing significant declines resulting in their at-risk status in Canada. We assessed intra-specific genetic variation, adaptive divergence, inbreeding, and genetic load across populations with different demographic histories using an annotated chromosome-scale reference genome and 66 whole-genome sequences. We found high genetic diversity and nine phylogenomic lineages across the continent with adaptive diversification of genes, but also high genetic load among lineages. We found highly divergent levels of inbreeding across individuals, including the loss of alleles by drift but not increased purging in inbred individuals, which had more homozygous deleterious alleles. We also found comparable frequencies of homozygous deleterious alleles between lineages regardless of nucleotide diversity. Thus, further inbreeding may need to be mitigated through conservation efforts. Our results highlight the \"double-edged sword\" of genetic diversity that may be representative of other species atrisk affected by anthropogenic activities.

Timing and quantity of sleep depend on a circadian (∼24-h) rhythm and a specific sleep requirement.1 Sleep curtailment results in a homeostatic rebound of more and deeper sleep, the latter reflected in increased electroencephalographic (EEG) slow-wave activity (SWA) during non-rapid eye movement (NREM) sleep.2 Circadian rhythms are synchronized by the light-dark cycle but persist under constant conditions.3,4,5 Strikingly, arctic reindeer behavior is arrhythmic during the solstices.6 Moreover, the Arctic\'s extreme seasonal environmental changes cause large variations in overall activity and food intake.7 We hypothesized that the maintenance of optimal functioning under these extremely fluctuating conditions would require adaptations not only in daily activity patterns but also in the homeostatic regulation of sleep. We studied sleep using non-invasive EEG in four Eurasian tundra reindeer (Rangifer tarandus tarandus) in Tromsø, Norway (69°N) during the fall equinox and both solstices. As expected, sleep-wake rhythms paralleled daily activity distribution, and sleep deprivation resulted in a homeostatic rebound in all seasons. Yet, these sleep rebounds were smaller in summer and fall than in winter. Surprisingly, SWA decreased not only during NREM sleep but also during rumination. Quantitative modeling revealed that sleep pressure decayed at similar rates during the two behavioral states. Finally, reindeer spent less time in NREM sleep the more they ruminated. These results suggest that they can sleep during rumination. The ability to reduce sleep need during rumination-undisturbed phases for both sleep recovery and digestion-might allow for near-constant feeding in the arctic summer.

Conservation breeding programs are increasingly used as recovery actions for wild animals; bringing founders into captivity to rear captive populations for future reintroduction into the wild. The International Union for the Conservation of Nature recommends that founders should come from genetically close populations and should have sufficient genetic diversity to avoid mating among relatives. Genomic data are highly informative for evaluating founders due to their high resolution and ability to capture adaptive divergence, yet, their application in that context remains limited. Woodland caribou are federally listed as a Species at Risk in Canada, with several populations facing extirpation, such as those in the Rocky Mountains of Alberta and British Columbia (BC). To prevent local extirpation, Jasper National Park (JNP) is proposing a conservation breeding program. We examined single nucleotide polymorphisms for 144 caribou from 11 populations encompassing a 200,0002 km area surrounding JNP to provide information useful for identifying appropriate founders for this program. We found that this area likely hosts a caribou metapopulation historically characterized by high levels of gene flow, which indicates that multiple sources of founders would be appropriate for initiating a breeding program. However, population structure and adaptive divergence analyses indicate that JNP caribou are closest to populations in the BC Columbia range, which also have suitable genetic diversity for conservation breeding. We suggest that collaboration among jurisdictions would be beneficial to implement the program to promote recovery of JNP caribou and possibly other caribou populations in the surrounding area, which is strategically at the periphery of the distribution of this endangered species.
UNASSIGNED: The online version contains supplementary material available at 10.1007/s10592-023-01540-3.

Assessing the effects of industrial development on wildlife is a key objective of managers and conservation practitioners. However, wildlife responses are often only investigated with respect to the footprint of infrastructure, even though human activity can strongly mediate development impacts. In Arctic Alaska, there is substantial interest in expanding energy development, raising concerns about the potential effects on barren-ground caribou (Rangifer tarandus granti). While caribou generally avoid industrial infrastructure, little is known about the role of human activity in moderating their responses, and whether managing activity levels could minimize development effects. To address this uncertainty, we examined the influence of traffic volume on caribou summer space use and road crossings in the Central Arctic Herd within the Kuparuk and Milne Point oil fields on the North Slope of Alaska. We first modeled spatiotemporal variation in hourly traffic volumes across the road system from traffic counter data using gradient-boosted regression trees. We then used generalized additive models to estimate nonlinear step selection functions and road-crossing probabilities from collared female caribou during the post-calving and insect harassment seasons, when they primarily interact with roads. Step selection analyses revealed that caribou selected areas further from roads (~1-3 km) during the post-calving and mosquito seasons and selected areas with lower traffic volumes during all seasons, with selection probabilities peaking when traffic was <5 vehicles/h. Using road-crossing models, we found that caribou were less likely to cross roads during the insect seasons as traffic increased, but that response dissipated as insect harassment became more severe. Past studies suggested that caribou exhibit behavioral responses when traffic exceeds 15 vehicles/h, but our results demonstrate behavioral responses at much lower traffic levels. Our results illustrate that vehicle activity mediates caribou responses to road infrastructure, information that can be used in future land-use planning to minimize the behavioral responses of caribou to industrial development in sensitive Arctic landscapes.