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Long-distance migrations are a striking, and strikingly successful, adaptation for highly mobile terrestrial animals in seasonal environments. However, it remains an open question whether migratory animals are more resilient or less resilient to rapidly changing environments. Furthermore, the mechanisms by which animals adapt or modify their migrations are poorly understood. We describe a dramatic shift of over 500 km in the wintering range of the Western Arctic Herd, a large caribou (Rangifer tarandus) herd in northwestern Alaska, an area that is undergoing some of the most rapid warming on Earth. Between 2012 and 2020, caribou switched from reliably wintering in maritime tundra in the southwesternmost portion of their range to more frequently wintering in mountainous areas to the east. Analysis of this range shift, in conjunction with nearly 200 documented mortality events, revealed that it was both broadly adaptive and likely driven by collective memory of poor winter conditions. Before the range shift, overwinter survival in the maritime tundra was high, routinely surpassing 95%, but falling to around 80% even as fewer animals wintered there. Meanwhile, in the increasingly used mountainous portion of the range, survival was intermediate and less variable across years compared to the extremes in the southern winter ranges. Thus, the shift only imperfectly mitigated overall increased mortality rates. The range shift has also been accompanied by changes in seasonal patterns of survival that are consistent with poorer nutritional intake in winter. Unexpectedly, the strongest single predictor of an individual's probability of migrating south was the overall survival of animals in the south in the preceding winter, suggesting that the range shift is in part driven by collective memory. Our results demonstrate the importance and use of collective decision making and memory for a highly mobile species for improving fitness outcomes in a dynamic, changing environment., (Published 2024. This article is a U.S. Government work and is in the public domain in the USA. Global Change Biology published by John Wiley & Sons Ltd.)

For prey, movement synchrony represents a potent antipredator strategy. Prey, however, must balance the costs and benefits of using conspecifics to mediate risk. Thus, the emergent patterns of risk-driven sociality depend on variation in space and in the predators and prey themselves. We applied the concept of predator-prey habitat domain, the space in which animals acquire food resources, to test the conditions under which individuals synchronize their movements relative to predator and prey habitat domains. We tested the response of movement synchrony of prey to predator-prey domains in two populations of ungulates that vary in their gregariousness and predator community: (i) elk, which are preyed on by wolves; and (ii) caribou, which are preyed on by coyotes and black bears. Prey in both communities responded to cursorial predators by increasing synchrony during seasons of greater predation pressure. Elk moved more synchronously in the wolf habitat domain during winter and caribou moved more synchronously in the coyote habitat domains during spring. In the winter, caribou increased movement synchrony when coyote and caribou domains overlapped. By integrating habitat domains with movement ecology, we provide a compelling argument for social behaviours and collective movement as an antipredator response. This article is part of the theme issue 'The spatial-social interface: A theoretical and empirical integration'.

Site fidelity-the tendency to reuse familiar spaces-is expected to improve fitness. Familiarity with the local environment is particularly crucial when resource demands or predation risk are high. Consequently, site fidelity often peaks during reproduction when energetic costs are high and offspring are vulnerable. For many species, the environment they experience is not solely a function of geography but also of the social environment. Social fidelity, the selection for familiar social environments, could constitute an independent or parallel strategy to spatial fidelity when considering behaviour at the spatial-social interface. Using global positioning system locations from caribou across Newfoundland, we tested whether females selected calving sites based on proximity to familiar conspecifics, in addition to geographical (spatial) fidelity. These strategies were synergistic, not alternative, and correlated across the population but more variable within individuals. We also tested whether either form of fidelity affected reproductive success. We failed to detect an effect of spatial or social fidelity on reproductive success in this population. Nevertheless, given the association between social and spatial fidelity and the demonstrated fitness consequences of site fidelity in other systems, familiar conspecifics and the potential benefits these social partners provide may be an underappreciated component driving site fidelity.This article is part of the theme issue 'The spatial-social interface: a theoretical and empirical integration'.

Encounters between animals occur when animals are close in space and time. Encounters are important in many ecological processes including sociality, predation and disease transmission. Despite this, there is little theory regarding the spatial distribution of encounters and no formal framework to relate environmental characteristics to encounters. The probability of encounter could be estimated with resource selection functions (RSFs) by comparing locations where encounters occurred to available locations where they may have occurred, but this estimate is complicated by the hierarchical nature of habitat selection. We developed a method to relate resources to the relative probability of encounter based on a scale-integrated habitat selection framework. This framework integrates habitat selection at multiple scales to obtain an appropriate estimate of availability for encounters. Using this approach, we related encounter probabilities to landscape resources. The RSFs describe habitat associations at four scales, home ranges within the study area, areas of overlap within home ranges, locations within areas of overlap, and encounters compared to other locations, which can be combined into a single scale-integrated RSF. We apply this method to intraspecific encounter data from two species: white-tailed deer (Odocoileus virginianus) and elk (Cervus elaphus) and interspecific encounter data from a two-species system of caribou (Rangifer tarandus) and coyote (Canis latrans). Our method produced scale-integrated RSFs that represented the relative probability of encounter. The predicted spatial distribution of encounters obtained based on this scale-integrated approach produced distributions that more accurately predicted novel encounters than a naïve approach or any individual scale alone. Our results highlight the importance of accounting for the conditional nature of habitat selection in estimating the habitat associations of animal encounters as opposed to 'naïve' comparisons of encounter locations with general availability. This method has direct relevance for testing hypotheses about the relationship between habitat and social or predator-prey behaviour and generating spatial predictions of encounters. Such spatial predictions may be vital for understanding the distribution of encounters driving disease transmission, predation rates and other population and community-level processes., (© 2024 The Author(s). Journal of Animal Ecology © 2024 British Ecological Society.)

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., (© 2024 The Author(s). Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Habitat loss is affecting many species, including the southern mountain caribou (Rangifer tarandus caribou) population in western North America. Over the last half century, this threatened caribou population's range and abundance have dramatically contracted. An integrated population model was used to analyze 51 years (1973-2023) of demographic data from 40 southern mountain caribou subpopulations to assess the effectiveness of population-based recovery actions at increasing population growth. Reducing potential limiting factors on threatened caribou populations offered a rare opportunity to identify the causes of decline and assess methods of recovery. Southern mountain caribou abundance declined by 51% between 1991 and 2023, and 37% of subpopulations were functionally extirpated. Wolf reduction was the only recovery action that consistently increased population growth when applied in isolation, and combinations of wolf reductions with maternal penning or supplemental feeding provided rapid growth but were applied to only four subpopulations. As of 2023, recovery actions have increased the abundance of southern mountain caribou by 52%, compared to a simulation with no interventions. When predation pressure was reduced, rapid population growth was observed, even under contemporary climate change and high levels of habitat loss. Unless predation is reduced, caribou subpopulations will continue to be extirpated well before habitat conservation and restoration can become effective., (© 2024 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Density dependence is a fundamental ecological process. In particular, animal habitat selection and social behavior often affect fitness in a density-dependent manner. The Ideal Free Distribution (IFD) and niche variation hypothesis (NVH) present distinct predictions associated with Optimal Foraging Theory about how the effect of habitat selection on fitness varies with population density. Using caribou (Rangifer tarandus) in Canada as a model system, we test competing hypotheses about how habitat specialization, social behavior, and annual reproductive success (co)vary across a population density gradient. Within a behavioral reaction norm framework, we estimate repeatability, behavioral plasticity, and covariance among social behavior and habitat selection to investigate the adaptive value of sociality and habitat selection. In support of NVH, but not the IFD, we find that at high density habitat specialists had higher annual reproductive success than generalists, but were also less social than generalists, suggesting the possibility that specialists were less social to avoid competition. Our study supports niche variation as a mechanism for density-dependent habitat specialization., (© 2024. The Author(s).)

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., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Inc. All rights reserved.)

AbstractUngulates can respond to changes in food supply by altering foraging behavior, digestive function, and metabolism. A multifaceted response to an environmental change is considered robust. Short seasons of plant growth make herbivores sensitive to changes in food supply because maintenance and production must be accomplished in less time with fewer options in a more fragile response. Caribou live at high latitudes where short summers constrain their response to changes in food supply. We measured the ability of female caribou to resist and tolerate changes in the quality and quantity of their food supply during winter and summer. Caribou resisted changes in food abundance and quality by changing food intake and physical activity with changes in daily temperature within each season. Peak food intake rose by 134% from winter pregnancy to summer lactation (98 vs. 229 g kg -0.75 d -1 ), as digestible requirements to maintain the body increased by 85% for energy (1,164 vs. 2,155 kJ kg -0.75 d -1 ) and by 266% for N (0.79 vs. 2.89 g N kg -0.75 d -1 ). Caribou required a diet with a digestible content of 12 kJ g -1 and 0.8% N in pregnancy, 18 kJ g -1 and 1.9% N in early lactation, and 11 kJ g -1 and 1.2% N in late lactation, which corresponds with the phenology of the wild diet. Female caribou tolerated restriction of ad lib. food intake to 58% of their energy requirement (680 vs. 1,164 kJ kg -0.75 d -1 ) during winter pregnancy and to 84% of their energy requirement (1,814 vs. 2,155 kJ kg -0.75 d -1 ) during summer lactation without a change in stress level, as indicated by fecal corticosterone concentration. Conversely, caribou can respond to increased availability of food with a spare capacity to process digestible energy and N at 123% (2,642 vs. 2,155 kJ kg -0.75 d -1 ) and 145% (4.20 vs. 2.89 g N kg -0.75 d -1 ) of those respective requirements during lactation. Robust responses to changes in food supply allow caribou to sustain reproduction, which would buffer demographic response. However, herds may decline when thresholds of behavioral resistance and physiological tolerance are frequently exceeded. Therefore, the challenge for managing declining populations of caribou and other robust species is to identify declines in robustness before their response becomes fragile.

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., (Published 2023. This article is a U.S. Government work and is in the public domain in the USA. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Muskox (Ovibos moschatus) and caribou (Rangifer tarandus groenlandicus) are wild ruminants that inhabit the Greenland tundra. They are part of the diet of many Greenlanders, being important sources of protein and micronutrients such as iron. The objective of this study is to analyse the element profiles of three tissues from these species: skeletal muscle, liver and adipose tissue, and to determine if they are affected by species and sex (male vs. female). Samples were obtained from annual hunts in two different regions of West Greenland. Element profiles were analysed using inductively-coupled plasma-optical emission spectrometry. The interaction between species and sex was only detected in Na (sodium) in the muscle and adipose tissue, where male and female caribou had the highest concentrations, respectively. The effect of sex was not statistically significant in the liver samples and only occasionally in the other tissues. Species was the most relevant factor in element profiles found in this study. Caribou had higher concentrations of K (potassium) and S (sulphur) in the muscle and liver. Fewer differences were detected between species in the adipose tissue, compared to the other tissues. These differences may reflect the feeding behaviour and the geographical location of both species. This study contributes to evaluate the element composition of the edible tissue of these wild ungulate species, as well as evaluating the factors of sex and species that could differentiate their composition., (© 2023. The Author(s).)

This study addresses for the first time the seasonal (winter-summer) dynamics of endogenous ethanol (EE) and acetaldehyde (EA) blood levels as well as of alcohol dehydrogenase (ADH) and aldehyde dehydrogenase (ALDH) activities and catalytic properties in the liver of the reindeer. EE blood level was 1.89 ± 0.11 mM in summer and 1.41 ± 0.10 mM (i.e. 1.34 times lower) in winter (p< 0.05). Seasonal dynamics of EA blood levels was opposite: 3.06 ± 0.28 μM in summer and 12.62 ± 0.76 μM (i.e. 4.12 times higher) in winter (p< 0.05). In winter, there was a decrease in activities of ADHII in the acetaldehyde reduction reaction (4.0 times) and ALDH (3.3 times) in the liver (p< 0.05). This led to an increased winter blood level of EA in cold-adapted animals. Our results suggest that EE and EA, as interrelated metabolites, make up one of the systems that regulate the metabolic rate and eventually reduce the intensity of energy metabolism in the reindeer. This consi derably extends the possibilities for northern species to exist under extreme environmental conditions of cold regions. Apparently, a major ph ysiological function of the system that includes EE, EA and relevant enzymes is to increase cold tolerance of an organism due to the inhibitory effect of EA on mitochondrial terminal oxidation which reduces the intensity of aerobic processes. [ABSTRACT FROM AUTHOR]

During summer and early autumn, reindeer and caribou (Rangifer tarandus) must exceed daily maintenance requirements for protein and energy to replenish body reserves for winter survival and reproduction. We estimated the maintenance requirements and the incremental costs of mass gain of captive female Siberian reindeer (R. t. tarandus) during summer and autumn by a mass balance approach. Intake (88 versus 56 g kg-0.75 day-1) and digestibility (89% versus 81%) of dry matter were greater in summer than in autumn on the same diet. Body mass increased over the summer and remained stable into autumn. At zero mass gain, reindeer required 911 kJ kg-0.75 day-1 digestible energy and 1.12 g kg-0.75 day-1 digestible nitrogen (N). Requirements of N were affected by endogenous losses of N in the feces that were 72-82% of total fecal N. During summer, reindeer would need to consume forage containing more than 9.5 ± 0.4 (SE) kJ g-1 digestible energy and 1.17 ± 0.4 (SE) % digestible N to gain body mass at intakes of 98 g kg-0.75 day-1. Reindeer could use a mix of graminoids and browse to meet energy demands and gain fat (6-18 kJ g-1) but may not be able to maintain or gain body protein in late summer as forages senesce and N contents decline below 1% of dry matter. [ABSTRACT FROM AUTHOR]

At temperate latitudes, the annual cycle of day length synchronizes circannual rhythms, and, in mammals, this is mediated via nocturnal production of the pineal hormone melatonin, proportional to the length of the night. Here, we studied circannual synchronization in an arctic species, the reindeer (Rangifer tarandus tarandus), which ceases to produce a rhythmic melatonin signal when it is exposed to extended periods of continuousmidwinter darkness and continuousmidsummer light. Using food intake, antler growth and moult as endpoints, we demonstrate that when animals living at 70°N are transferred from natural photoperiods in late autumn to either continuous light or continuous darkness, they undergo a conspicuous acceleration of the circannual programme. We conclude that rhythmical melatonin secretion, recommencing when the Sun reappears late in January, is required for proper timing of spring physiological responses, through a delaying effect on the circannual programme set in motion during the preceding autumn. [ABSTRACT FROM AUTHOR]

This longitudinal observational study was conducted to investigate the spontaneous effect of Giardia and Cryptosporidium infections on acute phase response (APR) in reindeer calves ( Rangifer tarandus tarandus ) in Finnish Lapland. Serum (n = 609) and faecal samples (n = 366) were collected from 54 reindeer calves aged zero to 33 days. The samples were analysed for Giardia , Cryptosporidium , acute phase proteins (APP) and γ-globulins. Linear regression models were used to investigate associations of early Giardia infection (before 12 days of life) with the response of APPs and acquiring of passive immunity. Giardia was detected in 100% and Cryptosporidium in 23% of calves. There was a negative association between early Giardia infection and γ-globulin concentrations (p = 0.032) and a positive association with serum amyloid A (SAA) concentrations (p = 0.042). The results suggest a protective effect of colostrum against Giardia infection and that early infection may induce activation of APR. [ABSTRACT FROM AUTHOR]

Reindeer in the arctic region live under very harsh conditions and may face temperatures below 233 K. Therefore, efficient conservation of body heat and water is important for their survival. Alongside their insulating fur, the reindeer nasal mechanism for heat and mass exchange during respiration plays a fundamental role. We present a dynamic model to describe the heat and mass transport that takes place inside the reindeer nose, where we account for the complicated geometrical structure of the subsystems that are part of the nose. The model correctly captures the trend in experimental data for the temperature, heat and water recovery in the reindeer nose during respiration. As a reference case, we model a nose with a simple cylindrical-like geometry, where the total volume and contact area are the same as those determined in the reindeer nose. A comparison of the reindeer nose with the reference case shows that the nose geometry has a large influence on the velocity, temperature and water content of the air inside the nose. For all investigated cases, we find that the total entropy production during a breathing cycle is lower for the reindeer nose than for the reference case. The same trend is observed for the total energy consumption. The reduction in the total entropy production caused by the complicated geometry is higher (up to -20%) at more extreme ambient conditions, when energy efficiency is presumably more important for the maintenance of energy balance in the animal. In the literature, a hypothesis has been proposed, which states that the most energy-efficient design of a system is characterized by equipartition of the entropy production. In agreement with this hypothesis, we find that the local entropy production during a breathing cycle is significantly more uniform for the reindeer nose than for the reference case. This suggests that natural selection has favored designs that give uniform entropy production when energy efficiency is an issue. Animals living in the harsh arctic climate, such as the reindeer, can therefore serve as inspiration for a novel industrial design with increased efficiency. [ABSTRACT FROM AUTHOR]

AbstractHeat waves are becoming more frequent across the globe and may impose severe thermoregulatory challenges for endotherms. Heat stress can induce both behavioral and physiological responses, which may result in energy deficits with potential fitness consequences. We studied the responses of reindeer ( Rangifer tarandus tarandus ), a cold-adapted ungulate, to a record-breaking heat wave in northern Finland. Activity, heart rate, subcutaneous body temperature, and body mass data were collected for 14 adult females. The post-heat wave autumn body masses were then analyzed against longitudinal body mass records for the herd from 1990 to 2021. With increasing air temperature during the day, reindeer became less active and had reduced heart rate and increased body temperature, reflecting both behavioral and physiological responses to heat stress. Although they increased activity in the late afternoon, they failed to compensate for lost foraging time on the hottest days (daily mean temperature ≥20°C), and total time active was reduced by 9%. After the heat wave, the mean September body mass of herd females ( 69.7 ± 6.6 kg, n = 52 ) was on average 16.4% ± 4.8% lower than predicted ( 83.4 ± 6.0 kg). Among focal females, individuals with the lowest levels of activity during the heat wave had the greatest mass loss during summer. We show how heat waves impose a thermoregulatory challenge on endotherms, resulting in mass loss, potentially as a result of the loss of foraging time. While it is well known that environmental conditions affect large herbivore fitness indirectly through decreased forage quality and limited water supply, direct effects of heat may be increasingly common in a warming climate.

Biodiversity is declining globally in response to multiple human stressors, including climate forcing. Nonetheless, local diversity trends are inconsistent in some taxa, obscuring contributions of local processes to global patterns. Arctic tundra diversity, including plants, fungi, and lichens, declined during a 15-year experiment that combined warming with exclusion of large herbivores known to influence tundra vegetation composition. Tundra diversity declined regardless of experimental treatment, as background growing season temperatures rose with sea ice loss. However, diversity declined slower with large herbivores than without them. This difference was associated with an increase in effective diversity of large herbivores as formerly abundant caribou declined and muskoxen increased. Efforts that promote herbivore diversity, such as rewilding, may help mitigate impacts of warming on tundra diversity.

In seasonal environments, animals should be adapted to match important life-history traits to when environmental conditions are optimal. Most animal populations therefore reproduce when resource abundance is highest to increase annual reproductive success. When facing variable, and changing, environments animals can display behavioural plasticity to acclimate to changing conditions. Behaviours can further be repeatable. For example, timing of behaviours and life history traits such as timing of reproduction may indicate phenotypic variation. Such variation may buffer animal populations against the consequences of variation and change. Our goal was to quantify plasticity and repeatability in migration and parturition timing in response to timing of snowmelt and green-up in a migratory herbivore (caribou, Rangifer tarandus, n = 132 ID-years) and their effect on reproductive success. We used behavioural reaction norms to quantify repeatability in timing of migration and timing of parturition in caribou and their plasticity to timing of spring events, while also quantifying phenotypic covariance between behavioural and life-history traits. Timing of migration for individual caribou was positively correlated with timing of snowmelt. The timing of parturition for individual caribou varied as a function of inter-annual variation in timing of snowmelt and green-up. Repeatability for migration timing was moderate, but low for timing of parturition. Plasticity did not affect reproductive success. We also did not detect any evidence of phenotypic covariance among any traits examined-timing of migration was not correlated with timing of parturition, and neither was there a correlation in the plasticity of these traits. Repeatability in migration timing suggests the possibility that the timing of migration in migratory herbivores could evolve if the repeatability detected in this study has a genetic or otherwise heritable basis, but observed plasticity may obviate the need for an evolutionary response. Our results also suggest that observed shifts in caribou parturition timing are due to plasticity as opposed to an evolutionary response to changing conditions. While this provides some evidence that populations may be buffered from the consequences of climate change via plasticity, a lack of repeatability in parturition timing could impede adaptation as warming increases., (© 2023 The Authors. Journal of Animal Ecology © 2023 British Ecological Society.)

Fragmentation of the boreal forest by linear features, including seismic lines, has destabilized predator-prey dynamics, resulting in the decline of woodland caribou (Rangifer tarandus caribou) populations. Restoration of human-altered habitat has therefore been identified as a critical management tool for achieving self-sustaining woodland caribou populations. However, only recently has testing of the response of caribou and other wildlife to restoration activities been conducted. Early work has centered around assessing changes in wildlife use of restored seismic lines. We evaluated whether restoration reduces the movement rates of predators and their associated prey, which is expected to decrease predator hunting efficiency and ultimately reduce caribou mortality. We developed a new method for using cameras to measure fine-scale movement by measuring speed as animals traveled between cameras in an array. We used our method to quantify speed of caribou, moose (Alces alces), bears (Ursus americanus), and wolves (Canis lupus) on treated (restored) and untreated seismic lines. Restoration treatments reduced travel speeds along seismic lines of wolves by 1.38 km/h, bears by 0.55 km/h, and caribou by 1.57 km/h, but did not reduce moose travel speeds. Reduced predator and caribou speeds on treated seismic lines are predicted to decrease encounter rates between predators and caribou and thus lower caribou kill rates. However, further work is needed to determine whether reduced movement rates result in reduced encounter rates with prey, and ultimately reduced caribou mortality., (© 2022 The Authors. Conservation Biology published by Wiley Periodicals LLC on behalf of Society for Conservation Biology.)

Based on the results of the analysis of nucleotide sequence polymorphism in the hypervariable fragment (left domain) of the mtDNA control region (D loop), the genetic diversity of wild reindeer from the eastern part of the Kola Peninsula (Murmansk oblast, Terskii region) was studied. Low values of the genetic diversity indices for Eurasian wild reindeer were detected. The effect of domestic reindeer breeding on the development of genetic diversity of the wild reindeer population in the studied region (indicating a low degree of hybridization of domesticated and wild reindeer) was estimated. [ABSTRACT FROM AUTHOR]

Abstract: Phylogenetic analyses provide information that can be useful in the conservation of genetic variation by identifying intraspecific genetic structure. Reconstruction of phylogenetic relationships requires the use of markers with the appropriate amount of variation relative to the timeframe and purpose of the study. Here, genetic structure and clustering are inferred from comparative analyses of three widely used mitochondrial markers, the CR, cytb and the COI region, merged and separately, using Eurasian reindeer as a model. A Bayesian phylogeny and a MJ network, both based on the merged dataset, indicate several distinct maternal haplotype clusters within Eurasian reindeer. In addition to confirm previously described clusters, two new subclusters were found. When comparing the results from the merged dataset with the results from analyses of the three markers separately, similar clustering was found in the CR and COI phylogenies, whereas the cytb region showed poor resolution. Phylogenetic analyses of the merged dataset and the CR revealed congruent results, implying that single sequencing analysis of the CR is an applicable method for studying the haplotype structure in Eurasian reindeer. [ABSTRACT FROM AUTHOR]

The nursing of non-offspring is referred to as allonursing. Reciprocity is a hypothesized cause of allonursing, but previous studies have not strongly supported or found no evidence in support for this hypothesis. Biological market theory was applied to 25 reindeer ( Rangifer tarandus) does and their calves to investigate allonursing reciprocity across bouts and within dyads and assess the diversity of allonursing partners. We also investigated whether variation in allonursing would be associated to relatedness within dyads. We recorded both the occurrence and the duration of 1027 successful allonursing solicitations. All does allonursed, and only three of the 25 calves were not observed being allonursed. Throughout the study, 234 allonursing dyadic pairs were observed. Does allonursed the calves of several does, but allonursing was not distributed evenly among all potential partners. Twenty does were members of at least 1 dyad with a high degree of reciprocity based on the number of allonursing bouts exchanged within the dyad. We found evidence of both relative and absolute allonursing reciprocity at the group level. Across bouts and within dyads, the reciprocal allonursing indexes varied greatly both for frequency and for duration, with an average tendency towards unidirectionality. Evidence of strong reciprocity within dyads was found in 32 dyads for number of allonursing bouts and in 25 dyads for duration of allonursing bouts. Across bouts and within dyads, the number of allonursing bouts received was not influenced by relatedness, allonursing bouts given or absolute rank difference. Allonursing was not interchanged for rank-related benefits. Our results provide evidence of reciprocal allonursing at the group level, across bouts and within dyads, and reciprocal allonursing among chosen partners. Our results point to the usefulness of applying the biological market theory to allonursing and of considering allonursing as a tradable commodity, traded for itself, among chosen partners. [ABSTRACT FROM AUTHOR]

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Arctic ecosystems are changing dramatically with warmer and wetter conditions resulting in complex interactions between herbivores and their forage. We investigated how Svalbard reindeer (Rangifer tarandus platyrhynchus) modify their late winter diets in response to long-term trends and interannual variation in forage availability and accessibility. By reconstructing their diets and foraging niches over a 17-year period (1995-2012) using serum δ 13 C and δ 15 N values, we found strong support for a temporal increase in the proportions of graminoids in the diets with a concurrent decline in the contributions of mosses. This dietary shift corresponds with graminoid abundance increases in the region and was associated with increases in population density, warmer summer temperatures and more frequent rain-on-snow (ROS) in winter. In addition, the variance in isotopic niche positions, breadths, and overlaps also supported a temporal shift in the foraging niche and a dietary response to extreme ROS events. Our long-term study highlights the mechanisms by which winter and summer climate changes cascade through vegetation shifts and herbivore population dynamics to alter the foraging niche of Svalbard reindeer. Although it has been anticipated that climate changes in the Svalbard region of the Arctic would be detrimental to this unique ungulate, our study suggests that environmental change is in a phase where conditions are improving for this subspecies at the northernmost edge of the Rangifer distribution., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Background: Dual-energy digital radiography (DEDR) has been shown to be a potential method to determine bone mineral density (BMD) and predict maximal load with similar accuracy as standard bone densitometry using DXA (dual-energy X-ray absorptiometry). In addition to bone density, bone geometry has also been shown to have effect on bone fragility and fracture risk. Purpose: To examine the combination of BMD and geometry parameters, as determined from a DEDR experiment, to predict bone maximal load. Material and Methods: Reindeer femora (n = 47) were imaged at two energies (79 kVp and 100 kVp) using a clinical digital radiography system. BMD was determined in four regions from these images using the DXA calculation principle. Various geometrical parameters were determined from the 79 kVp image. Femora were mechanically tested using axial loading configuration. Pearson correlation coefficients were determined between geometrical parameters and BMDs or maximal load. Multiple stepwise linear regression analysis was used to find the best combination to predict bone maximal load. Results: From the geometrical parameters, femoral shaft diameter (FSD) and femoral neck axis length (FNAL) correlated best with the maximal load (r = 0.629 and r = 0.446, P < 0.01, respectively). The best combination of parameters to predict bone fragility was BMD at Ward's triangle, FSD and FNAL (r = 0.787, P < 0.05), whereas the correlation coefficient between BMD at Ward's triangle and maximal load was 0.653 (P < 0.05). Conclusion: The combination of DEDR-based BMD and geometrical parameters predicts reindeer bone maximal load with reasonable accuracy and the combined analysis improves the prediction of maximal load compared to BMD prediction only. [ABSTRACT FROM AUTHOR]

Abstract: Energy deprivation during the wintertime poses a challenge to skeletal muscle function. The objective of the present study was to determine the expression of specific calcium (Ca2+) regulating proteins in skeletal muscle of the reindeer during wintertime adaptation and moderate weight loss. To assess the possible changes in the dihydropyridine receptor (Cacna1), ryanodine receptor (RyR1), sarco(endo)plasmic reticulum Ca2+-ATPase (SERCA1) and calsequestrin 1 (CASQ1) proteins, six mature female reindeer were held under experimental outdoor conditions with limited energy intake and muscle biopsies were taken from m. semitendinosus. In addition, muscle biopsies from m. semitendinosus and m. gluteus medius taken from freely ranging reindeer after slaughter (October, n =7; January, n =7) were included in the study. The muscle samples were analysed by Western blot. The animals lost 10–21% of their body mass (BM) during the restricted feeding, with no change in the muscle total protein content. In general, the concentrations of nearly all Ca2+ regulating components measured were maintained as a response to restricted feeding in experimental and freely ranging reindeer. Interestingly, the expression of Cacna1 was 2.3-fold up-regulated after the restricted feeding period in the m. semitendinosus of the experimental reindeer. On the contrary, in freely ranging animals, the expression of SERCA1 and RyR1 decreased 3.5- and 3.2-fold, respectively, in January compared to October. To conclude, the expression of Ca2+ regulating proteins is not altered during moderate weight loss and winter adaptation in order to maintain effective muscle contraction capacity. Up-regulation of Cacna1, however, indicates the ability to allocate Ca2+ cycling capacity to both oxidative and glycolytic metabolic pathways during times of energy deprivation. [Copyright &y& Elsevier]

Recovering endangered species is a difficult and often controversial task that challenges status quo land uses. Southern Mountain caribou are a threatened ecotype of caribou that historically ranged in southwestern Canada and northwestern USA and epitomize the tension between resource extraction, biodiversity conservation, and Indigenous Peoples' treaty rights. Human-induced habitat alteration is considered the ultimate cause of caribou population declines, whereby an increased abundance of primary prey-such as moose and deer-elevates predator populations and creates unsustainable caribou mortality. Here we focus on the Klinse-Za and Quintette subpopulations, part of the endangered Central Group of Southern Mountain caribou in British Columbia. These subpopulations were trending toward immediate extirpation until a collaborative group initiated recovery by implementing two short-term recovery actions. We test the effectiveness of these recovery actions-maternity penning of adult females and their calves, and the reduction of a primary predator, wolves-in increasing vital rates and population growth. Klinse-Za received both recovery actions, whereas Quintette only received wolf reductions, providing an opportunity to test efficacy between recovery actions. Between 1995 and 2021, we followed 162 collared female caribou for 414 animal-years to estimate survival and used aerial counts to estimate population abundance and calf recruitment. We combined these data in an integrated population model to estimate female population growth, total population abundance, and recovery action effectiveness. Results suggest that the subpopulations were declining rapidly (λ = 0.90-0.93) before interventions and would have been functionally extirpated (<10 animals) within 10-15 years. Wolf reduction increased population growth rates by ~0.12 for each subpopulation. Wolf reduction halted the decline of Quintette caribou and allowed them to increase (λ = 1.05), but alone would have only stabilized the Klinse-Za (λ = 1.02). However, maternity penning in the Klinse-Za increased population growth by a further ~0.06, which when combined with wolf reductions, allowed populations to grow (λ = 1.08). Taken together, the recovery actions in these subpopulations increased adult female survival, calf recruitment, and overall population growth, more than doubling abundance. Our results suggest that maternity penning and wolf reductions can be effective at increasing caribou numbers in the short term, while long-term commitments to habitat protection and restoration are made., (© 2022 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Indigenous Peoples around the northern hemisphere have long relied on caribou for subsistence and for ceremonial and community purposes. Unfortunately, despite recovery efforts by federal and provincial agencies, caribou are currently in decline in many areas across Canada. In response to recent and dramatic declines of mountain caribou populations within their traditional territory, West Moberly First Nations and Saulteau First Nations (collectively, the "Nations") came together to create a new vision for caribou recovery on the lands they have long stewarded and shared. The Nations focused on the Klinse-Za subpopulation, which had once encompassed so many caribou that West Moberly Elders remarked that they were "like bugs on the landscape." The Klinse-Za caribou declined from ~250 in the 1990s to only 38 in 2013, rendering Indigenous harvest of caribou nonviable and infringing on treaty rights to a subsistence livelihood. In collaboration with many groups and governments, this Indigenous-led conservation initiative paired short-term population recovery actions, predator reduction and maternal penning, with long-term habitat protection in an effort to create a self-sustaining caribou population. Here, we review these recovery actions and the promising evidence that the abundance of Klinse-Za caribou has more than doubled from 38 animals in 2013 to 101 in 2021, representing rapid population growth in response to recovery actions. With looming extirpation averted, the Nations focused efforts on securing a landmark conservation agreement in 2020 that protects caribou habitat over a 7986-km 2 area. The Agreement provides habitat protection for >85% of the Klinse-Za subpopulation (up from only 1.8% protected pre-conservation agreement) and affords moderate protection for neighboring caribou subpopulations (29%-47% of subpopulation areas, up from 0%-20%). This Indigenous-led conservation initiative has set both the Indigenous and Canadian governments on the path to recover the Klinse-Za subpopulation and reinstate a culturally meaningful caribou hunt. This effort highlights how Indigenous governance and leadership can be the catalyst needed to establish meaningful conservation actions, enhance endangered species recovery, and honor cultural connections to now imperiled wildlife., (© 2022 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

In 2004 to 2006, studies on free-ranging reindeer and musk oxen on Wrangel island were performed to estimate the nutrient value, daily intake, and digestibility of forage in different seasons of the year. Forage digestibility was determined from the ratio between the contents of indigestible components (lignin) in forage and animal feces. Daily forage intake was calculated from data on daily feces excretion and forage digestibility. The amount of feces and parameters of life activity of individual animals were estimated by following their 24-hour tracks. The results show that the daily intake of forage (dry weight) in musk oxen amounts to 7.8 kg in the summer period and decreases to 6.1 kg in the snow period (March-April). Grazing reindeer in the snow period consume 3.9 kg of forage per day. Forage digestibility in reindeer reaches 56% in March to April and decreases to 52% in June. In musk oxen, forage digestibility in different seasons (March, June, September) varies within a similar range (53-57%). Nutrition parameters (diet composition, forage intake rate and digestibility) of reindeer and musk oxen on grazing grounds of Wrangel Island are similar. Metabolizable energy consumption in both animals during the winter period is at the maintenance level (917-930 kJ/kg0.75 body weight). In the summer period, this parameter in musk oxen increases to 1163 kJ/kg0.75 body weight to meet the energy demand of the animals. [ABSTRACT FROM AUTHOR]

Reindeer are numerous and widespread across the northern Holarctic. They are efficient long distance migrants and are able to cope with variations in substrate, such as ice, snow, uneven forest floor, wetland and flat grassland. However, as with the vast majority of quadrupedal vertebrates, no quantitative musculoskeletal anatomical information exists for these animals making it difficult to analyse the biomechanics of their locomotor behaviour. In this paper, we describe the gross anatomy of the limb musculature and quantify muscle and tendon morphology. Reindeer show slight hindlimb dominance in muscle and tendon mass, with muscle mass primarily proximally situated and tendon distally situated. Extensor muscles are heavier than flexors, but tendon mass is broadly similar in both extensors and flexors. The only complete quadrupedal data sets available for comparison are for hares and greyhounds making it difficult to identify general patterns. There are no obvious body mass effects and reindeer often comes out as intermediate between hare and greyhound. However, greyhound seem less hindlimb dominated in terms of muscle but both greyhound and hare have much higher masses of tendon compared to reindeer, particularly in their hindlimbs. All these quadrupeds show the commonly observed trait of much larger tendons and less massive muscles in distal limb segments; this reduces the inertial cost of accelerating the limbs. Generally, there is a dearth of available quantitative anatomical data of complete animals. This lack of information is hindering attempts to gain a better understanding of musculoskeletal function in quadrupeds. [ABSTRACT FROM AUTHOR]

Abstract: The deer ked (Lipoptena cervi) is a haematophagous parasitic fly of cervids that spread to Finland in the early 1960''s. Presently its northern distribution limit lies at approximately 65°N and it is gradually spreading northwards. In Finland the principal host species has been the moose (Alces alces), but the deer ked is about to establish contact with another potential host, the semi-domesticated reindeer (Rangifer tarandus tarandus) causing possible threats to reindeer health and management. The aim of this study was to investigate if the deer ked would have an influence on the welfare of the reindeer. Eighteen adult reindeer were divided into three experimental groups: the control group and two infected groups with 300 deer keds per reindeer introduced in August–September. One of the infected groups was treated with subcutaneous ivermectin in November. To gather comprehensive data on potential health hazards caused by the deer ked a wide array of physiological variables was measured during and at the end of the experiment in December. The keds caused no clear changes in the complete blood count, plasma clinical chemistry, amino acids, endocrinology, energy stores, enzyme activities or tissue fatty acid profiles of the host. The haematological, clinical chemical and endocrinological values displayed changes that could be related to the seasonal physiological adaptations of the species. In conclusion, at the duration and intensity of infection that were employed, the effects of the deer ked on the measured physiological variables of the reindeer were insignificant. [Copyright &y& Elsevier]

Abstract: Ruminant species differ in the degree that their rumen contents are stratified but are similar insofar that only very fine particles are passed from the forestomach to the lower digestive tract. We investigated the passage kinetics of fluid and particle markers (2, 10 and 20mm) in fistulated cattle (Bos primigenius f. taurus), muskoxen (Ovibos moschatus), reindeer (Rangifer tarandus) and moose (Alces alces) on different diets. The distribution of dry matter in the rumen and the viscosity of rumen fluids suggested that the rumen contents were more stratified in muskoxen than moose. Correspondingly, as in previous studies, the species differed in the ratio of mean retention times of small particles to fluids in the reticulorumen, which was highest in cattle (2.03) and muskoxen (1.97–1.98), intermediate in reindeer (1.70) and lowest in moose (0.98–1.29). However, the ratio of large to small particle retention did not differ between the species, indicating similarity in the efficiency of the particle sorting mechanism. Passage kinetics of the two largest particle classes did not differ, indicating that particle retention is not a continuous function of particle size but rather threshold-dependent. Overall, the results suggest that fluid flow through the forestomach differs between ruminant species. A lower relative fluid passage, such as in moose, might limit species to a browse-based dietary niche, whereas a higher relative fluid passage broadens the dietary niche options and facilitates the inclusion of, or specialization on, grass. The function of fluid flow in the ruminant forestomach should be further investigated. [Copyright &y& Elsevier]

Abstract: In young ruminants, the reticular groove ensures that ingested milk is channelled past the forestomach to avoid malfermentation. It has been speculated that some adult wild ruminants, in particular browsing species, maintain a functional oesophageal (reticular) groove, that soluble nutrients can thus bypass the rumen, and that thus the energetic gain from the diet can be increased. We inserted a fluid marker (Co–EDTA) via cannula into the rumen and simultaneously fed a diet that contained a second fluid marker (Sm–EDTA), and analysed the faecal marker excretion patterns, in muskoxen (Ovibos moschatus, n =4 in two experiments each), reindeer (Rangifer tarandus, n =4 in a total of six experiments) and moose (Alces alces, n =1 in one experiment). In no case was the orally fed marker excreted distinctively earlier than the marker inserted into the rumen, which indicates that substantial bypass did not occur in these animals. However, differences between the three species in the excretion of the two markers from the rumen are consistent with hypothetical differences in the stratification of rumen contents. We suggest that effects previously ascribed to a “rumen bypass” in wild ruminants most likely reflect differences in the passage from the rumen. [Copyright &y& Elsevier]

Transport of 22Na and 14C-butyrate across the ruminal epithelium of captive reindeer fed a concentrate diet in summer (n=5) and in winter (n=5) and from free-ranging reindeer taken from summer (n=3) and winter pasture (n=5) was measured in vitro in Ussing chambers. Significant amounts of both Na+ and butyrate were transported across the isolated epithelium without any external driving force. The ruminal transport of Na+ and butyrate were interacting, as evidenced by both the observed amiloride-induced reduction of net butyrate-transport and by the positive correlation between net transport of butyrate and Na+. Amiloride also reduced the net transport of Na+ without significantly affecting the short-circuit current, indicating the presence of an apical Na+/H+ exchanger in the ruminal epithelium of reindeer. The captive reindeer increased the dry matter intake of a constant quality concentrate from winter to summer, but this neither affected their ruminal transport capacity nor their ruminal surface enlargement factor (SEF). Free-ranging reindeer increased their ruminal transport capacity for Na+ and butyrate from summer to winter but simultaneously reduced their ruminal SEF. The present data indicate that this food-induced increase in transport capacity was attributed to changes in the nutrient composition of the diet. [ABSTRACT FROM AUTHOR]

Studies the variation in lactate transport in red blood cells of racing species like reindeer, horses, humans and dogs. Total lactate-transport activity at different lactate concentrations; Role of different transporters with inhibitors; Effect of training on lactate-transport activity among the species.

This study tested the hypothesis that the annual cycle in heart rate (HR) in reindeer is, at least in part, a consequence of seasonal fluctuation in voluntary-food intake. Heart rate and daily dry matter voluntary-food intake (DDMVFI) were recorded in two captive female reindeer (Rangifer tarandus tarandus) from April 1995 to August 1996. Heart rate was measured continuously in each animal for 20–24 h for 7 days each month using Polar® Sport Testers (PST); DDMVFI was measured in each animal daily for 17 months. Modal daily heart rate (MDHR) and DDMVFI fluctuated seasonally in close synchrony, both reaching maxima in July and minima in January. The relationship between HR and DDMVFI was investigated experimentally by manipulating the level of feeding in a stepwise manner in May, when appetite was low and in August, when DDMVFI was close to maximum. Heart rate showed stepwise changes in close synchrony with the changes in levels of feeding. These results suggest that the seasonal increase in HR in summer is a consequence of increased food intake and, likewise, decreased HR in winter is a consequence of reduced food intake. The observed relationship between food intake and HR presumably reflects changes in cardiac output and/or the rate of flow of blood to the gastrointestinal tract which are influenced by meal size. [ABSTRACT FROM AUTHOR]

Climate change will lead to more frequent and more severe fires in some areas of boreal forests, affecting the distribution and availability of late-successional forest communities. These forest communities help to protect globally significant carbon reserves beneath permafrost layers and provide habitat for many animal species, including forest-dwelling caribou. Many caribou populations are declining, yet the mechanisms by which changing fire regimes could affect caribou declines are poorly understood. We analyzed resource selection of 686 GPS-collared female caribou from three ecotypes and 15 populations in a ~600,000 km 2 region of northwest Canada and eastern Alaska. These populations span a wide gradient of fire frequency but experience low levels of human-caused habitat disturbance. We used a mixed-effects modeling framework to characterize caribou resource selection in response to burns at different seasons and spatiotemporal scales, and to test for functional responses in resource selection to burn availability. We also tested mechanisms driving observed selection patterns using burn severity and lichen cover data. Caribou avoided burns more strongly during winter relative to summer and at larger spatiotemporal scales relative to smaller scales. During the winter, caribou consistently avoided burns at both spatiotemporal scales as burn availability increased, indicating little evidence of a functional response. However, they decreased their avoidance of burns during summer as burn availability increased. Burn availability explained more variation in caribou selection for burns than ecotype. Within burns, caribou strongly avoided severely burned areas in winter, and this avoidance lasted nearly 30 years after a fire. Caribou within burns also selected higher cover of terrestrial lichen (an important caribou food source). We found a negative relationship between burn severity and lichen cover, confirming that caribou avoidance of burns was consistent with lower lichen abundance. Consistent winter avoidance of burns and severely burned areas suggests that caribou will experience increasing winter habitat loss as fire frequency and severity increase. Our results highlight the potential for climate-induced alteration of natural disturbance regimes to affect boreal biodiversity through habitat loss. We suggest that management strategies prioritizing protection of core winter range habitat with lower burn probabilities would provide important climate-change refugia for caribou., (© 2022 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Describes exercise-induced changes in blood composition of racing reindeer. Resting values of hematological data; Protein and amino acid concentrations in serum.

Copyright of Blut: Zeitschrift für die Gesamte Blutforschung is the property of Springer Nature and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Background: The alternative prey hypothesis describes the mechanism for apparent competition whereby the mortality of the secondary prey species increases (and population size decreases decreases) by the increased predation by the shared predator if the population size of the primary prey decreases. Apparent competition is a process where the abundance of two co-existing prey species are negatively associated because they share a mutual predator, which negatively affects the abundance of both prey Here, we examined whether alternative prey and/or apparent competition hypothesis can explain the population dynamics and reproductive output of the secondary prey, wild forest reindeer (Rangifer tarandus fennicus) in Finland, in a predator-prey community in which moose (Alces alces) is the primary prey and the wolf (Canis lupus) is the generalist predator., Methods: We examined a 22-year time series (1996-2017) to determine how the population size and the calf/female ratio of wild forest reindeer in Eastern Finland were related to the abundances of wolf and moose. Only moose population size was regulated by hunting. Summer predation of wolves on reindeer focuses on calves. We used least squares regression (GLS) models (for handling autocorrelated error structures and resulting pseudo-R2s) and generalized linear mixed (GLMs) models (for avoidance of negative predictions) to determine the relationships between abundances. We performed linear and general linear models for the calf/female ratio of reindeer., Results and Synthesis: The trends in reindeer population size and moose abundance were almost identical: an increase during the first years and then a decrease until the last years of our study period. Wolf population size in turn did not show long-term trends. Change in reindeer population size between consecutive winters was related positively to the calf/female ratio. The calf/female ratio was negatively related to wolf population size, but the reindeer population size was related to the wolf population only when moose abundance was entered as another independent variable. The wolf population was not related to moose abundance even though it is likely to consist the majority of the prey biomass. Because reindeer and moose populations were positively associated, our results seemed to support the alternative prey hypothesis more than the apparent competition hypothesis. However, these two hypotheses are not mutually exclusive and the primary mechanism is difficult to distinguish as the system is heavily managed by moose hunting. The recovery of wild forest reindeer in eastern Finland probably requires ecosystem management involving both habitat restoration and control of species abundances., Competing Interests: The authors have declared that no competing interests exist.

Rangifer tarandus has experienced recent drastic population size reductions throughout its circumpolar distribution and preserving the species implies genetic diversity conservation. To facilitate genomic studies of the species populations, we improved the genome assembly by combining long read and linked read and obtained a new highly accurate and contiguous genome assembly made of 13,994 scaffolds (L90 = 131 scaffolds). Using de novo transcriptome assembly of RNA-sequencing reads and similarity with annotated human gene sequences, 17,394 robust gene models were identified. As copy number variations (CNVs) likely play a role in adaptation, we additionally investigated these variations among 20 genomes representing three caribou ecotypes (migratory, boreal and mountain). A total of 1,698 large CNVs (length > 1 kb) showing a genome distribution including hotspots were identified. 43 large CNVs were particularly distinctive of the migratory and sedentary ecotypes and included genes annotated for functions likely related to the expected adaptations. This work includes the first publicly available annotation of the caribou genome and the first assembly allowing genome architecture analyses, including the likely adaptive CNVs reported here., (© 2021 Prunier et al.)

As global climate change progresses, wildlife management will benefit from knowledge of demographic responses to climatic variation, particularly for species already endangered by other stressors. In Canada, climate change is expected to increasingly impact populations of threatened woodland caribou (Rangifer tarandus caribou) and much focus has been placed on how a warming climate has potentially facilitated the northward expansion of apparent competitors and novel predators. Climate change, however, may also exert more direct effects on caribou populations that are not mediated by predation. These effects include meteorological changes that influence resource availability and energy expenditure. Research on other ungulates suggests that climatic variation may have minimal impact on low-density populations such as woodland caribou because per-capita resources may remain sufficient even in "bad" years. We evaluated this prediction using demographic data from 21 populations in western Canada that were monitored for various intervals between 1994 and 2015. We specifically assessed whether juvenile recruitment and adult female survival were correlated with annual variation in meteorological metrics and plant phenology. Against expectations, we found that both vital rates appeared to be influenced by annual climatic variation. Juvenile recruitment was primarily correlated with variation in phenological conditions in the year prior to birth. Adult female survival was more strongly correlated with meteorological conditions and declined during colder, more variable winters. These responses may be influenced by the life history of woodland caribou, which reside in low-productivity refugia where small climatic changes may result in changes to resources that are sufficient to elicit strong demographic effects. Across all models, explained variation in vital rates was low, suggesting that other factors had greater influence on caribou demography. Nonetheless, given the declining trajectories of many woodland caribou populations, our results highlight the increased relevance of recovery actions when adverse climatic conditions are likely to negatively affect caribou demography., Competing Interests: The authors have declared that no competing interests exist.

Seasonal energetic challenges may constrain an animal's ability to respond to changing individual and environmental conditions. Here, we investigated variation in heart rate, a well-established proxy for metabolic rate, in Svalbard reindeer ( Rangifer tarandus platyrhynchus ), a species with strong seasonal changes in foraging and metabolic activity. In 19 adult females, we recorded heart rate, subcutaneous temperature and activity using biologgers. Mean heart rate more than doubled from winter to summer. Typical drivers of energy expenditure, such as reproduction and activity, explained a relatively limited amount of variation (2-6% in winter and 16-24% in summer) compared to seasonality, which explained 75% of annual variation in heart rate. The relationship between heart rate and subcutaneous temperature depended on individual state via body mass, age and reproductive status, and the results suggested that peripheral heterothermy is an important pathway of energy management in both winter and summer. While the seasonal plasticity in energetics makes Svalbard reindeer well-adapted to their highly seasonal environment, intraseasonal constraints on modulation of their heart rate may limit their ability to respond to severe environmental change. This study emphasizes the importance of encompassing individual state and seasonal context when studying energetics in free-living animals. This article is part of the theme issue 'Measuring physiology in free-living animals (Part II)'.

The present review provides a comprehensive assessment on the basic aspects of the reproduction of the Rangifer male with a special focus on pubertal development, and the seasonal dynamics of gametogenesis, androgenesis, antler cycle and other physiological and behavioural characteristics of the rutting male. A greater understanding of the reproductive physiology of the males is needed to improve existing management strategies of semi-domestic stocks in the changing climate and to develop species-specific assisted reproductive technologies for improving the in situ and ex situ conservation programs to preserve endangered subspecies as well as to facilitate the transfer of genetic resources between fragmented wild or semi-domestic populations., (Copyright © 2021 The Authors. Published by Elsevier B.V. All rights reserved.)

Movement provides a link between individual behavioral ecology and the spatial and temporal variation in an individual's landscape. Individual variation in movement traits is an important axis of animal personality, particularly in the context of foraging ecology. We tested whether individual caribou ( Rangifer tarandus ) displayed plasticity in movement and space-use behavior across a gradient of resource aggregation. We quantified first-passage time and range-use ratio as proxies for movement-related foraging behavior and examined how these traits varied at the individual level across a foraging resource gradient. Our results suggest that individuals adjusted first-passage time but not range-use ratio to maximize access to high-quality foraging resources. First-passage time was repeatable, and intercepts for first-passage time and range-use ratio were negatively correlated. Individuals matched first-passage time but not range-use ratio to the expectations of our patch-use model that maximized access to foraging resources, a result that suggests that individuals acclimated their movement patterns to accommodate both intra- and interannual variation in foraging resources on the landscape. Collectively, we highlight repeatable movement and space-use tactics and provide insight into how individual plasticity in movement interacts with landscape processes to affect the distribution of behavioral phenotypes and potentially fitness and population dynamics.

Reindeer (Rangifer tarandus) have evolved elaborate nasal turbinate structures that are perfused via a complex vascular network. These are subject to thermoregulatory control, shifting between heat conservation and dissipation, according to the animal's needs. The three-dimensional design of the turbinate structures is essential in the sense that they determine the efficiency with which heat and water are transferred between the structure and the respired air. The turbinates have already a relatively large surface area at birth, but the structures have yet not reached the complexity of the mature animal. The aim of this study was to elucidate the structure-function relationship of the heat exchange process. We have used morphometric and physiological data from newborn reindeer calves to construct a thermodynamic model for respiratory heat and water exchange and present novel results for the simulated respiratory energy losses of calves in the cold. While the mature reindeer effectively conserves heat and water through nasal counter-current heat exchange, the nose of the calf has not yet attained a similar efficiency. We speculate that this is probably related to structure-size limitations and more favourable climate conditions during early life. The fully developed structure-function relationship may serve as inspiration for engineering design. Simulations of different extents of mucosal vascularization suggest that the abundance and pattern of perfusion of veins in the reindeer nasal mucosa may contribute to the control of temperature profiles, such that nasal cavity tissue is sufficiently warm, but not excessively so, keeping heat dissipation within limits.

Reindeer are semi-domesticated ruminants that have adapted to the challenging northern Eurasian environment characterized by long winters and marked annual fluctuations in daylight. We explored the genetic makeup behind their unique characteristics by de novo sequencing the genome of a male reindeer and conducted gene family analyses with nine other mammalian species. We performed a population genomics study of 23 additional reindeer representing both domestic and wild populations and several ecotypes from various geographic locations. We assembled 2.66 Gb (N50 scaffold of 5 Mb) of the estimated 2.92 Gb reindeer genome, comprising 27,332 genes. The results from the demographic history analysis suggested marked changes in the effective population size of reindeer during the Pleistocene period. We detected 160 reindeer-specific and expanded genes, of which zinc finger proteins (n = 42) and olfactory receptors (n = 13) were the most abundant. Comparative genome analyses revealed several genes that may have promoted the adaptation of reindeer, such as those involved in recombination and speciation (PRDM9), vitamin D metabolism (TRPV5, TRPV6), retinal development (PRDM1, OPN4B), circadian rhythm (GRIA1), immunity (CXCR1, CXCR2, CXCR4, IFNW1), tolerance to cold-triggered pain (SCN11A) and antler development (SILT2). The majority of these characteristic reindeer genes have been reported for the first time here. Moreover, our population genomics analysis suggested at least two independent reindeer domestication events with genetic lineages originating from different refugial regions after the Last Glacial Maximum. Taken together, our study has provided new insights into the domestication, evolution and adaptation of reindeer and has promoted novel genomic research of reindeer.