Winter condition variability decreases the economic sustainability of reindeer husbandry.

Wild and semidomesticated reindeer are one of the key species in Arctic and subarctic areas, and their population dynamics are closely tied to winter conditions. Difficult snow conditions have been found to decrease the calving success and survivability of reindeer, but the economic effects of variation in winter conditions on reindeer husbandry have not been studied. In this study, we combine state-of-the-art economic-ecological modeling with the analysis of annual reindeer management reports from Finland. These contain local knowledge of herding communities. We quantify the occurrence probabilities of different types of winters from annual management reports and analyze the effects of this variation in winter conditions on reindeer husbandry using an age- and sex-structured bioeconomic reindeer-lichen model. Our results show that difficult winters decrease the net revenues of reindeer husbandry. However, they also protect lichen pastures from grazing, thereby increasing future net revenues. Nonetheless, our solutions show that the variability of winter conditions overall decrease the net income of herders compared to constant winter conditions. Low lichen biomass appears to make reindeer management more sensitive to the effects of difficult winter conditions. We also found that it is economically sensible to use supplementary feeding during difficult winters, but the net revenues still decrease compared to average winters because of the high feeding costs. Overall, our analysis suggests that the increasing variability of winter conditions due to climate change will decrease net revenues in reindeer husbandry. This decrease will still occur even if the most extreme effects of climate change do not occur. This study shows that combining a state-of-the-art bioeconomic model and practitioner knowledge can bring compatible insights, ideas, results, and a bottom-up perspective to the discussion.

Fluctuating optimum and temporally variable selection on breeding date in birds and mammals.

Temporal variation in natural selection is predicted to strongly impact the evolution and demography of natural populations, with consequences for the rate of adaptation, evolution of plasticity, and extinction risk. Most of the theory underlying these predictions assumes a moving optimum phenotype, with predictions expressed in terms of the temporal variance and autocorrelation of this optimum. However, empirical studies seldom estimate patterns of fluctuations of an optimum phenotype, precluding further progress in connecting theory with observations. To bridge this gap, we assess the evidence for temporal variation in selection on breeding date by modeling a fitness function with a fluctuating optimum, across 39 populations of 21 wild animals, one of the largest compilations of long-term datasets with individual measurements of trait and fitness components. We find compelling evidence for fluctuations in the fitness function, causing temporal variation in the magnitude, but not the direction of selection. However, fluctuations of the optimum phenotype need not directly translate into variation in selection gradients, because their impact can be buffered by partial tracking of the optimum by the mean phenotype. Analyzing individuals that reproduce in consecutive years, we find that plastic changes track movements of the optimum phenotype across years, especially in bird species, reducing temporal variation in directional selection. This suggests that phenological plasticity has evolved to cope with fluctuations in the optimum, despite their currently modest contribution to variation in selection.

Early-life conditions determine the between-individual heterogeneity in plasticity of calving date in reindeer.

Phenotypic plasticity has become a key concept to enhance our ability to understand the adaptive potential of species to track the pace of climate change by allowing a relatively rapid adjustment of life-history traits. Recently, population-level trends of an earlier timing of reproduction to climate change have been highlighted in many taxa, but only few studies have explicitly taken into consideration between-individual heterogeneity in phenotypic plasticity. Using a long-term data of a semi-domesticated reindeer (Rangifer tarandus) population, we demonstrated that females differed greatly in their mean calving date but only slightly in the magnitude of their plastic response to the amount of precipitation in April. We also showed that despite the absence of a population trend, females individually responded to the amount of precipitation in April by delaying their calving dates. Females' calving date under average climatic conditions was best predicted by their birthdate, by their physical condition in March-April-May before their first calving season and by their first calving date. The degree of their phenotypic plasticity was not dependent on any of the females' attributes early in life tested in this study. However, females who delayed their calving dates in response to a higher amount of precipitation in April slightly produced less calves over their reproductive life. These findings confirmed that early-life conditions of female reindeer can shape their phenotypic value during reproductive life, supporting the importance of maternal effects in shaping individuals' lifetime reproductive success. Whether females differed in the magnitude of their plastic response to climatic changes has received contrasted responses for various ungulate species. This calls for more research to enhance our understanding of the underlying mechanisms, leading to the complexity of plastic responses among populations to cope with current climate change.

Response of reindeer mating time to climatic variability.

BACKGROUND: The breeding time of many species has changed over the past 2-3 decades in response to climate change. Yet it is a key reproductive trait that affects individual's parturition time and reproductive success, and thereby population dynamics. In order to predict how climate change will affect species' viability, it is crucial to understand how species base their reproductive efforts on environmental cues. RESULTS: By using long-term datasets of mating behaviours and copulation dates recorded since 1996 on a semi-domesticated reindeer population, we showed that mating time occurred earlier in response to weather conditions at different key periods in their annual breeding cycle. The mating time occurred earlier following a reducing snow cover in early spring, colder minimum temperatures in the last 2 weeks of July and less precipitation in August-September. CONCLUSIONS: The mediated effect of a reduced snow cover in early spring on improving individuals' pre-rut body weight through a better availability of late winter food and reduced costs of locomotion on snow would explain that mating time has occurred earlier overtime. A lower level of insect harassment caused by colder maximum temperatures in July might have caused an advance in mating time. Less precipitation in August-September also caused the mating time to occur earlier, although the direct effects of the last two weather variables were not mediated through the pre-rut body weight of individuals. As such, the causal effects of weather conditions on seasonal timing of animals are still unclear and other mechanisms than just body weight might be involved (e.g. socio-biological factors). The plastic response of reindeer mating time to climatic variability, despite supplemental feeding occurring in late April, demonstrated that environmental factors may have a greater influence on reproductive outputs than previously thought in reindeer.

Modelling reindeer rut activity using on-animal acoustic recorders and machine learning.

For decades, researchers have employed sound to study the biology of wildlife, with the aim to better understand their ecology and behaviour. By utilizing on-animal recorders to capture audio from freely moving animals, scientists can decipher the vocalizations and glean insights into their behaviour and ecosystem dynamics through advanced signal processing. However, the laborious task of sorting through extensive audio recordings has been a major bottleneck. To expedite this process, researchers have turned to machine learning techniques, specifically neural networks, to streamline the analysis of data. Nevertheless, much of the existing research has focused predominantly on stationary recording devices, overlooking the potential benefits of employing on-animal recorders in conjunction with machine learning. To showcase the synergy of on-animal recorders and machine learning, we conducted a study at the Kutuharju research station in Kaamanen, Finland, where the vocalizations of rutting reindeer were recorded during their mating season. By attaching recorders to seven male reindeer during the rutting periods of 2019 and 2020, we trained convolutional neural networks to distinguish reindeer grunts with a 95% accuracy rate. This high level of accuracy allowed us to examine the reindeers' grunting behaviour, revealing patterns indicating that older, heavier males vocalized more compared to their younger, lighter counterparts. The success of this study underscores the potential of on-animal acoustic recorders coupled with machine learning techniques as powerful tools for wildlife research, hinting at their broader applications with further advancement and optimization.

A Summer Heat Wave Reduced Activity, Heart Rate, and Autumn Body Mass in a Cold-Adapted Ungulate.

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.

The onset in spring and the end in autumn of the thermal and vegetative growing season affect calving time and reproductive success in reindeer.

A developing trophic mismatch between the peak of energy demands by reproducing animals and the peak of forage availability has caused many species' reproductive success to decrease. The match-mismatch hypothesis (MMH) is an appealing concept that can be used to assess such fitness consequences. However, concerns have been raised on applying the MMH on capital breeders such as reindeer because the reliance on maternal capita rather than dietary income may mitigate negative effects of changing phenologies. Using a long-term dataset of reindeer calving dates recorded since 1970 in a semidomesticated reindeer population in Finnish Lapland and proxies of plant phenology; we tested the main hypothesis that the time lag between calving date and the plant phenology in autumn when females store nutrient reserves to finance reproduction would lead to consequences on reproductive success, as the time lag with spring conditions would. As predicted, the reproductive success of females of the Kutuharju reindeer population was affected by both the onset of spring green-up and vegetative senescence in autumn as calves were born heavier and with a higher first-summer survival when the onset of the vegetation growth was earlier and the end of the thermal growing season the previous year was earlier as well. Our results demonstrated that longer plant growing seasons might be detrimental to reindeer's reproductive success if a later end is accompanied by a reduced abundance of mushrooms.

Stabilizing selection and adaptive evolution in a combination of two traits in an arctic ungulate.

Stabilizing selection is thought to be common in wild populations and act as one of the main evolutionary mechanisms, which constrain phenotypic variation. When multiple traits interact to create a combined phenotype, correlational selection may be an important process driving adaptive evolution. Here, we report on phenotypic selection and evolutionary changes in two natal traits in a semidomestic population of reindeer (Rangifer tarandus) in northern Finland. The population has been closely monitored since 1969, and detailed data have been collected on individuals since they were born. Over the length of the study period (1969-2015), we found directional and stabilizing selection toward a combination of earlier birth date and heavier birth mass with an intermediate optimum along the major axis of the selection surface. In addition, we demonstrate significant changes in mean traits toward earlier birth date and heavier birth mass, with corresponding genetic changes in breeding values during the study period. Our results demonstrate evolutionary changes in a combination of two traits, which agree closely with estimated patterns of phenotypic selection. Knowledge of the selective surface for combinations of genetically correlated traits are vital to predict how population mean phenotypes and fitness are affected when environments change.

Winter and spring climatic conditions influence timing and synchrony of calving in reindeer.

In a context of climate change, a mismatch has been shown to occur between some species' reproductive phenology and their environment. So far, few studies have either documented temporal trends in calving phenology or assessed which climatic variables influence the calving phenology in ungulate species, yet the phenology of ungulates' births affects offspring survival and population's recruitment rate. Using a long-term dataset (45 years) of birth dates of a semi-domesticated reindeer population in Kaamanen, North Finland, we show that calving season has advanced by ~ 7 days between 1970 and 2016. Advanced birth dates were associated with lower precipitation and a reduced snow cover in April and warmer temperatures in April-May. Improved females' physical condition in late gestation due to warmer temperatures in April-May and reduced snow conditions in April probably accounted for such advance in calving date. On the other hand, a lengthening of the calving season was reported following a warmer temperature in January, a higher number of days when mean temperature exceeds 0°C in October-November and a decreasing snow cover from October to November. By affecting the inter-individual heterogeneity in the plastic response of females' calving date to better climatic conditions in fall and winter, climatic variability contributed to weaken the calving synchrony in this herd. Whether variability in climatic conditions form environmental cues for the adaptation of calving phenology by females to climate change is however uncertain, but it is likely. As such this study enhances our understanding on how reproductive phenology of ungulate species would be affected by climate change.

Male phenotypic quality influences offspring sex ratio in a polygynous ungulate.

Evolutionary models of sex ratio adjustment applied to mammals have ignored that females may gain indirect genetic benefits from their mates. The differential allocation hypothesis (DAH) predicts that females bias the sex ratio of their offspring towards (more costly) males when breeding with an attractive male. We manipulated the number of available males during rut in a polygynous ungulate species, the reindeer (Rangifer tarandus), and found that a doubling of average male mass (and thus male attractiveness) in the breeding herd increased the proportion of male offspring from approximately 40 to 60%. Paternity analysis revealed indeed that males of high phenotypic quality sired more males, consistent with the DAH. This insight has consequences for proper management of large mammal populations. Our study suggests that harvesting, by generating a high proportion of young, small and unattractive mates, affects the secondary sex ratio due to differential allocation effects in females. Sustainable management needs to consider not only the direct demographic changes due to harvest mortality and selection, but also the components related to behavioural ecology and opportunities for female choice.

Parameterization and validation of an ungulate-pasture model.

Ungulate grazing and trampling strongly affect pastures and ecosystems throughout the world. Ecological population models are used for studying these systems and determining the guidelines for sustainable and economically viable management. However, the effect of trampling and other resource wastage is either not taken into account or quantified with data in earlier models. Also, the ability of models to describe the herbivore impact on pastures is usually not validated. We used a detailed model and data to study the level of winter- and summertime lichen wastage by reindeer and the effects of wastage on population sizes and management. We also validated the model with respect to its ability of predicting changes in lichen biomass and compared the actual management in herding districts with model results. The modeling efficiency value (0.75) and visual comparison between the model predictions and data showed that the model was able to describe the changes in lichen pastures caused by reindeer grazing and trampling. At the current lichen biomass levels in the northernmost Finland, the lichen wastage varied from 0 to 1 times the lichen intake during winter and from 6 to 10 times the intake during summer. With a higher value for wastage, reindeer numbers and net revenues were lower in the economically optimal solutions. Higher wastage also favored the use of supplementary feeding in the optimal steady state. Actual reindeer numbers in the districts were higher than in the optimal steady-state solutions for the model in 18 herding districts out of 20. Synthesis and applications. We show that a complex model can be used for analyzing ungulate-pasture dynamics and sustainable management if the model is parameterized and validated for the system. Wastage levels caused by trampling and other causes should be quantified with data as they strongly affect the results and management recommendations. Summertime lichen wastage caused by reindeer is higher than expected, which suggests that seasonal pasture rotation should be used to prevent the heavy trampling of winter lichen pastures during summer. In the present situation, reindeer numbers in northernmost Finland are in most cases higher than in the management solutions given by the model.

Infestation by Hypoderma tarandi in reindeer calves from northern Finland--prevalence and risk factors.

Serum samples from 953 reindeer (Rangifer t. tarandus) calves-of-the-year from 21 reindeer herding co-operatives in Finland in slaughter season 2006/2007 were assayed for antibodies against Hypodermin C by an ELISA detecting IgG. Data on presence of Hypoderma tarandi larvae on 12,327 reindeer hides from 17 of the 21 herding co-operatives in slaughter season 2005/2006 were included for support. ELISA showed a seroprevalence of 60.9%, with no significant difference between females and males, and increase with latitude (southernmost and northernmost co-operatives examined, Pudasjärvi and Kaldoaivi, 11.8 and 100% of seropositives, respectively). The proportion of larva positive hides (range 0.5-60% per co-operative) was low compared to the proportion of seropositives. Also the proportion of larva positive hides increased with latitude. Our findings indicated that high latitude combined with open landscape, presence of low vegetation and high reindeer density provided more favorable conditions for sustaining of high degree of warble fly infestation, and furthermore, that any possible effect of ivermectin treatment on infestation rate was ruled out by the higher effect by the above factors.

Climate change promotes the emergence of serious disease outbreaks of filarioid nematodes.

Filarioid parasites represent major health hazards with important medical, veterinary, and economic implications, and considerable potential to affect the everyday lives of tens of millions of people globally (World Health Organization, 2007). Scenarios for climate change vary latitudinally and regionally and involve direct and indirect linkages for increasing temperature and the dissemination, amplification, and invasiveness of vector-borne parasites. High latitude regions are especially influenced by global climate change and thus may be prone to altered associations and dynamics for complex host-pathogen assemblages and emergence of disease with cascading effects on ecosystem structure. Although the potential for substantial ecological perturbation has been identified, few empirical observations have emanated from systems across the Holarctic. Coincidental with decades of warming, and anomalies of high temperature and humidity in the sub-Arctic region of Fennoscandia, the mosquito-borne filarioid nematode Setaria tundra is now associated with emerging epidemic disease resulting in substantial morbidity and mortality for reindeer and moose. We describe a host-parasite system that involves reindeer, arthropods, and nematodes, which may contribute as a factor to ongoing declines documented for this ungulate species across northern ecosystems. We demonstrate that mean summer temperatures exceeding 14 degrees C drive the emergence of disease due to S. tundra. An association between climate and emergence of filarioid parasites is a challenge to ecosystem services with direct effects on public health, sustainability of free-ranging and domestic ungulates, and ultimately food security for subsistence cultures at high latitudes.

Ecological role of reindeer summer browsing in the mountain birch (Betula pubescens ssp. czerepanovii) forests: effects on plant defense, litter decomposition, and soil nutrient cycling.

Mammalian herbivores commonly alter the concentrations of secondary compounds in plants and, by this mechanism, have indirect effects on litter decomposition and soil carbon and nutrient cycling. In northernmost Fennoscandia, the subarctic mountain birch (Betula pubescens ssp. czerepanovii) forests are important pasture for the semidomestic reindeer (Rangifer tarandus). In the summer ranges, mountain birches are intensively browsed, whereas in the winter ranges, reindeer feed on ground lichens, and the mountain birches remain intact. We analyzed the effect of summer browsing on the concentrations of secondary substances, litter decomposition, and soil nutrient pools in areas that had been separated as summer or winter ranges for at least 20 years, and we predicted that summer browsing may reduce levels of secondary compounds in the mountain birch and, by this mechanism, have an indirect effect on the decomposition of mountain birch leaf litter and soil nutrient cycling. The effect of browsing on the concentration of secondary substances in the mountain birch leaves varied between different years and management districts, but in some cases, the concentration of condensed tannins was lower in the summer than in the winter ranges. In a reciprocal litter decomposition trial, both litter origin and emplacement significantly affected the litter decomposition rate. Decomposition rates were faster for the litter originating from and placed into the summer range. Soil inorganic nitrogen (N) concentrations were higher in the summer than in the winter ranges, which indicates that reindeer summer browsing may enhance the soil nutrient cycling. There was a tight inverse relationship between soil N and foliar tannin concentrations in the winter range but not in the summer range. This suggests that in these strongly nutrient-limited ecosystems, soil N availability regulates the patterns of resource allocation to condensed tannins in the absence but not in the presence of browsing.

Reproductive success in reindeer males in a herd with varying sex ratio.

In polygynous species, male reproductive success is often correlated with dominance status of individual males and sex ratio in the population. Reindeer, Rangifer tarandus, is a polygynous species, and here we compared the variation in male reproductive success and dominance status during two successive years in a herd with a male:female sex ratio of 1:7 and 1:3. Copulations were recorded, together with data on male dominance hierarchy and size of mating groups. Male reproductive success was estimated by paternity analysis of calves using microsatellite DNA markers. The distribution of reproductive success among the males was highly skewed for both years with the most dominant male also being the most successful. The largest mating group was established in the herd with the least skew in sex ratio. In this herd some of the adult males present were less reproductively successful than some of the more subordinate younger males. Estimates of the mating group size of males, correcting for dominance status when more than one male is present in the groups, gave good prediction of individual males' reproductive success.