Decomposing demographic contributions to the effective population size with moose as a case study.

Levels of random genetic drift are influenced by demographic factors, such as mating system, sex ratio and age structure. The effective population size (Ne ) is a useful measure for quantifying genetic drift. Evaluating relative contributions of different demographic factors to Ne is therefore important to identify what makes a population vulnerable to loss of genetic variation. Until recently, models for estimating Ne have required many simplifying assumptions, making them unsuitable for this task. Here, using data from a small, harvested moose population, we demonstrate the use of a stochastic demographic framework allowing for fluctuations in both population size and age distribution to estimate and decompose the total demographic variance and hence the ratio of effective to total population size (Ne /N) into components originating from sex, age, survival and reproduction. We not only show which components contribute most to Ne /N currently, but also which components have the greatest potential for changing Ne /N. In this relatively long-lived polygynous system we show that Ne /N is most sensitive to the demographic variance of older males, and that both reproductive autocorrelations (i.e., a tendency for the same individuals to be successful several years in a row) and covariance between survival and reproduction contribute to decreasing Ne /N (increasing genetic drift). These conditions are common in nature and can be caused by common hunting strategies. Thus, the framework presented here has great potential to increase our understanding of the demographic processes that contribute to genetic drift and viability of populations, and to inform management decisions.

Opposing fitness consequences of habitat use in a harvested moose population.

Landscape changes are happening at an unprecedented pace, and together with high levels of wildlife harvesting humans have a large effect on wildlife populations. A thorough knowledge of their combined influence on individual fitness is important to understand factors affecting population dynamics. The goal of the study was to assess the individual consistency in the use of risky habitat types, and how habitat use was related to fitness components and life-history strategies. Using data from a closely monitored and harvested population of moose Alces alces, we examined how individual variation in offspring size, reproduction and survival was related to the use of open grasslands; a habitat type that offers high-quality forage during summer, but at the cost of being more exposed to hunters in autumn. The use of this habitat type may therefore involve a trade-off between high mortality risk and forage maximization. There was a high repeatability in habitat use, which suggests consistent behaviour within individuals. Offspring number and weight were positively related to the mothers' use of open grasslands, whereas the probability of surviving the subsequent harvest season was negatively related to the use of the same habitat type. As a consequence, we found a nonsignificant relationship between habitat use and lifetime fitness. The study suggests that harvesting, even if intended to be nonselective with regard to phenotypes, may be selective towards animals with specific behaviour and life-history strategies. As a consequence, harvesting can alter the life-history composition of the population and target life-history strategies that would be beneficial for individual fitness and population growth in the absence of hunting.

Effect of gamma radiation on the production of bystander signals from three earthworm species irradiated in vivo.

The effect of gamma radiation delivered over 24 h on the induction of bystander signals of three earthworm species exposed in vivo was investigated: A. chlorotica, A. caliginosa, and E. tetraedra. Worms were exposed to external gamma irradiation (Co-60 source) for 24 h and samples of head, body, and clitellum were dissected from exposed and control worms and placed in culture medium for 24 h at 19 C. The harvested medium was filtered and assayed for expression of bystander signals using both clonogenic and mitochondrial reporter assays. Different responses were observed in the different species and in the different tissues. A. chlorotica worm-treated reporters show insignificant mitochondrial response for all sections, yet a significant clonogenic reduction in survival for body sections. A. caliginosa worm-treated reporters show a significant mitochondrial response for some sections and insignificant mitochondrial response and insignificant reduction in clonogenic survival for the rest. E. tetraedra worms from a control site show significant evidence of bystander signalling, measured by mitochondrial response in reporter cells, for all sections while those harvested from a contaminated site show insignificant changes in baseline signalling when exposed to the challenge dose. In vivo exposure of earthworm species shows evidence of bystander signalling using two different reporter assays. This effect varied between the different species and tissues. There is also evidence of attenuated bystander signalling in worms harvested from a site contaminated with radiation.

Fitness correlates of age at primiparity in a hunted moose population.

Trade-offs between fitness-related traits are predicted from the principle of resource allocation, where increased fecundity or parental investment leads to reduced future reproduction or survival. However, fitness traits can also be positively correlated due to individual differences (e.g. body mass). Age at primiparity could potentially explain variation in individual fitness either because early primiparity is costly, or it may lead to higher lifetime reproductive success. Based on long-term monitoring and genetic parentage assignment of an island population of moose, we quantified reproductive performance and survival, and examined whether early maturing females have higher total calf production than late maturing females. We explored if harvesting of calves affected the subsequent reproductive success of their mothers, i.e. also due to a post-weaning cost of reproduction, and whether there are any intergenerational effects of female reproductive success. There was a positive relationship between current and future reproduction. The probability to reproduce was lower for females that were unsuccessful the year before, indicating a strong quality effect on productivity. Females that started to reproduce as 2-year olds had a slightly higher total calf production compared to those starting at age three or four. High-performing mothers were also correlated with daughters that performed well in terms of reproductive success. Our results suggest that the observed individual heterogeneity in fitness could be associated with differences in age at primiparity. This heterogeneity was not affected by reproductive costs associated with tending for a calf post-weaning.

Population properties affect inbreeding avoidance in moose.

Mechanisms reducing inbreeding are thought to have evolved owing to fitness costs of breeding with close relatives. In small and isolated populations, or populations with skewed age- or sex distributions, mate choice becomes limited, and inbreeding avoidance mechanisms ineffective. We used a unique individual-based dataset on moose from a small island in Norway to assess whether inbreeding avoidance was related to population structure and size, expecting inbreeding avoidance to be greater in years with larger populations and even adult sex ratios. The probability that a potential mating event was realized was negatively related to the inbreeding coefficient of the potential offspring, with a stronger relationship in years with a higher proportion or number of males in the population. Thus, adult sex ratio and population size affect the degree of inbreeding avoidance. Consequently, conservation managers should aim for sex ratios that facilitate inbreeding avoidance, especially in small and isolated populations.

Moose body mass variation revisited: disentangling effects of environmental conditions and genetics.

Large-scale geographical variation in phenotypic traits within species is often correlated to local environmental conditions and population density. Such phenotypic variation has recently been shown to also be influenced by genetic structuring of populations. In ungulates, large-scale geographical variation in phenotypic traits, such as body mass, has been related to environmental conditions and population density, but little is known about the genetic influences. Research on the genetic structure of moose suggests two distinct genetic lineages in Norway, structured along a north-south gradient. This corresponds with many environmental gradients, thus genetic structuring provides an additional factor affecting geographical phenotypic variation in Norwegian moose. We investigated if genetic structure explained geographical variation in body mass in Norwegian moose while accounting for environmental conditions, age and sex, and if it captured some of the variance in body mass that previously was attributed to environmental factors. Genetic structuring of moose was the most important variable in explaining the geographic variation in body mass within age and sex classes. Several environmental variables also had strong explanatory power, related to habitat diversity, environmental seasonality and winter harshness. The results suggest that environmental conditions, landscape characteristics, and genetic structure should be evaluated together when explaining large-scale patterns in phenotypic characters or life history traits. However, to better understand the role of genetic and environmental effects on phenotypic traits in moose, an extended individual-based study of variation in fitness-related characters is needed, preferably in an area of convergence between different genetic lineages.

Cold season dose rate contributions from gamma, radon, thoron or progeny in legacy mines with high natural background radiation.

In areas with high natural background radiation, underground cavities tend to have high levels of airborne radionuclides. Within mines, occupancy may involve significant exposure to airborne radionuclides like radon (222Rn), thoron (220Rn) and progeny. The Fen carbonatite complex in Norway has legacy mines going through bedrock with significantly elevated levels of uranium (238U) and especially thorium (232Th), and significant levels of their progeny 222Rn and 220Rn. There are also significantly elevated levels of gamma radiation in these mines. These mines are naturally chimney ventilated and release large volumes of air to the outdoors giving a large local outdoor impact. We placed alpha track detectors at several localities within these mines to measure airborne radionuclides and measured gamma radiation of bedrock at each locality. The bedrock within the mines shows levels up to 1900 Bq kg-1 for 238U, 12 000 Bq kg-1 for 232Th and gamma dose rates up to 11 µSv h-1. Maximum levels of airborne radionuclides were 45 000 Bq m-3 for 220Rn and 6900 Bq m-3 for 222Rn. In addition, we measured levels of thoron progeny (TnP). In order to estimate radiation dose contribution, TnP should be assessed rather than 220Rn, but deposition-based detectors may be biased by the airflow of mine-draft. We present dose rate contributions using UNSCEAR dose conversion factors, and correcting for airflow bias, finding a combined cold season dose rate within these mines of 17-24 µSv h-1. Interestingly, fractional dose rate contributions vary from 0.02 to 0.6 for gamma, 0.33 to 0.95 for radon and 0.1 to 0.25 for TnP.

Increased barium levels in recent marine sediments from the Norwegian and Barents Seas suggest impact of hydrocarbon drilling and production.

Barium (Ba) in recent marine sediments can originate from natural and anthropogenic sources including discharges from the oil and gas industry. In this study, we use data from the Norwegian and Barents Seas to assess whether Ba in recent marine sediments has increased due to these discharges. To account for Ba in detrital material, we normalise all samples with respect to aluminosilicate by calculating an enrichment factor. We use statistical modelling to control for parameters related to sedimentation. We present results that suggest increased Ba levels in recent sediments that coincide with the timing of hydrocarbon drilling and production. This is supported by geographical differences on a large scale that relate to proximity to hydrocarbon drilling and production. Among 243 sampling stations, we identify 73 locations exhibiting enrichment of Ba in the upper 6 of sediment. At these locations, Ba is 1.55 to 3.55 times higher than the levels that can be expected from the shale average when Ba in detrital matter is accounted for. Excess Ba is reported in sediment surface samples in areas important to fisheries like the Lofoten area and the western Barents Sea.

Tools for harmonized data collection at exposure situations with naturally occurring radioactive materials (NORM).

Naturally occurring radioactive materials (NORM) contribute to the dose arising from radiation exposure for workers, public and non-human biota in different working and environmental conditions. Within the EURATOM Horizon 2020 RadoNorm project, work is ongoing to identify NORM exposure situations and scenarios in European countries and to collect qualitative and quantitative data of relevance for radiation protection. The data obtained will contribute to improved understanding of the extent of activities involving NORM, radionuclide behaviours and the associated radiation exposure, and will provide an insight into related scientific, practical and regulatory challenges. The development of a tiered methodology for identification of NORM exposure situations and complementary tools to support uniform data collection were the first activities in the mentioned project NORM work. While NORM identification methodology is given in Michalik et al., 2023, in this paper, the main details of tools for NORM data collection are presented and they are made publicly available. The tools are a series of NORM registers in Microsoft Excel form, that have been comprehensively designed to help (a) identify the main NORM issues of radiation protection concern at given exposure situations, (b) gain an overview of materials involved (i.e., raw materials, products, by-products, residues, effluents), c) collect qualitative and quantitative data on NORM, and (d) characterise multiple hazards exposure scenarios and make further steps towards development of an integrated risk and exposure dose assessment for workers, public and non-human biota. Furthermore, the NORM registers ensure standardised and unified characterisation of NORM situations in a manner that supports and complements the effective management and regulatory control of NORM processes, products and wastes, and related exposures to natural radiation worldwide.

A methodology for the systematic identification of naturally occurring radioactive materials (NORM).

Naturally occurring radioactive materials (NORM) are present worldwide and under certain circumstances (e.g., human activities) may give radiation exposure to workers, local public or occasional visitors and non-human biota (NHB) of the surrounding ecosystems. This may occur during planned or existing exposure situations which, under current radiation protection standards, require identification, management, and regulatory control as for other practices associated with man-made radionuclides that may result in the exposure of people and NHB. However, knowledge gaps exist with respect to the extent of global and European NORM exposure situations and their exposure scenario characteristics, including information on the presence of other physical hazards, such as chemical and biological ones. One of the main reasons for this is the wide variety of industries, practices and situations that may utilise NORM. Additionally, the lack of a comprehensive methodology for identification of NORM exposure situations and the absence of tools to support a systematic characterisation and data collection at identified sites may also lead to a gap in knowledge. Within the EURATOM Horizon 2020 RadoNorm project, a methodology for systematic NORM exposure identification has been developed. The methodology, containing consecutive tiers, comprehensively covers situations where NORM may occur (i.e., minerals and raw materials deposits, industrial activities, industrial products and residues and their applications, waste, legacies), and thus, allows detailed investigation and complete identification of situations where NORM may present a radiation protection concern in a country. Details of the tiered methodology, with practical examples on harmonised data collection using a variety of existing sources of information to establish NORM inventories, are presented in this paper. This methodology is flexible and thus applicable to a diversity of situations. It is intended to be used to make NORM inventory starting from the scratch, however it can be used also to systematise and complete existing data.

Reduced soil fauna decomposition in a high background radiation area.

Decomposition of litter and organic matter is a very important soil ecosystem function where soil fauna play an important role. Knowledge of the responses in decomposition and soil fauna to different stressors is therefore crucial. However, the extent to which radioactivity may affect soil fauna is not so well known. There are some results showing effects on soil fauna at uranium mines and near Chernobyl from relatively high levels of anthropogenic radionuclides. We hypothesize that naturally occurring radionuclides affect soil fauna and thus litter decomposition, which will covary with radionuclide levels when accounting for important soil parameters. We have therefore used standardised litterbags with two different mesh sizes filled with birch leaves (Betula pubescens) to assess litter decomposition in an area with enhanced levels of naturally occurring radionuclides in the thorium (232Th) and uranium (238U) decay chains while controlling for variation in important soil parameters like pH, organic matter content, moisture and large grain size. We show that decomposition rate is higher in litterbags with large mesh size compared to litterbags with a fine mesh size that excludes soil fauna. We also find that litter dried at room temperature is decomposed at a faster rate than litter dried in oven (60°C). This was surprising given the associated denaturation of proteins and anticipated increased nutritional level but may be explained by the increased stiffness of oven-dried litter. This result is important since different studies often use either oven-dried or room temperature-dried litter. Taking the above into account, we explore statistical models to show large and expected effects of soil parameters but also significant effects on litter decomposition of the naturally occurring radionuclide levels. We use the ERICA tool to estimate total dose rate per coarse litterbag for four different model organisms, and in subsequent different statistical models we identify that the model including the dose rates of a small tube-shape is the best statistical model. In another statistical model including soil parameters and radionuclide distributions, 226Ra (or uranium precursory radionuclides) explain variation in litter decomposition while 228Ra (and precursors) do not. This may hint to chemical toxicity effects of uranium. However, when combining this model with the best model, the resulting simplified model is equal to the tube-shape dose-rate model. There is thus a need for more research on how naturally occurring radionuclides affect soil fauna, but the study at hand show the importance of an ecosystem approach and the ecosystem parameter soil decomposition.

Realism and usefulness of multispecies experiment designs with regard to application in radioecology: A review.

Multispecies experiments like microcosms and mesocosms are widely used in many fields of research but not in radioecology. In radioecology, size limitations are important as large experimental volumes involve problems with waste (radionuclides), or shielding, absorption and available space in gamma fields (often within a climate chamber). We have therefore performed a literature review (ISI Web of Science, n = 406) of the design and properties of multispecies effect studies <100 L in size and with three or more mentioned taxa in other research fields to assess their suitability to radioecology. Studies with more mentioned taxa assess structural ecosystem parameters more often than studies with fewer mentioned taxa, while the opposite trend is seen for indirect effects/interactions. Studies of indirect effects benefit from more replicates and longer experiments. Almost all studies assess some ecosystem level parameter but only a quarter take a holistic approach assessing both structural and functional as well as indirect effects. We find that most cosms are custom-made systems, rather than standardised designs. Unmanipulated cosms consist of excised portions of the natural environment with a higher number of mentioned taxa, high ecological complexity and high realism, but have a relatively low replicability. In contrast, standardised cosms with fewer mentioned taxa have less ecological complexity but much higher replicability. This literature review shows that smaller cosm sizes have similar ecological complexity (e.g. number of taxa and trophic levels) and experimental duration as larger sized cosms, allowing for ecologically-relevant investigations, despite their small size. We encourage multispecies radioecology studies, preferably with environmental relevant doses and sufficient detail on dosimetry.

Indoor and Outdoor Exposure to Radon, Thoron and Thoron Decay Products in a NORM Area with Highly Elevated Bedrock Thorium and Legacy Mines.

Radon (222Rn) and thoron (220Rn), and especially their short-lived decay products, are major contributors to dose received by the public from naturally occurring radioactive material (NORM), particularly in areas with elevated levels of naturally occurring radionuclides. Mining in such areas can involve ventilation of high amounts of these gases, which may influence outdoor levels. In this work, we assessed indoor and outdoor levels of 222Rn, 220Rn and 220Rn decay products (TnDP) in close proximity to an area with elevated bedrock levels of thorium (232Th) and a NORM legacy mining site with high natural ventilation. We assess municipal buildings at distances from a few hundred meters to 2 km from the NORM legacy mines. In some buildings, high indoor levels of 222Rn were observed in winter, as expected for temperate areas. In summer, high indoor levels of 222Rn and 220Rn were observed in some buildings, and very low associated levels of TnDP in actively ventilated buildings may suggest entry by ventilation and an outdoor source. Outdoor levels of TnDP increased with decreased distance from the legacy mines, suggesting dispersal from these during both summer and winter.

Norwegian monitoring (1990-2015) of the marine environment around the sunken nuclear submarine Komsomolets.

Norway has monitored the marine environment around the sunken Russian nuclear submarine Komsomolets since 1990. This study presents an overview of 25 years of Norwegian monitoring data (1990-2015). Komsomolets sank in 1989 at a depth of 1680 m in the Norwegian Sea while carrying two nuclear torpedoes in its armament. Subsequent Soviet and Russian expeditions to Komsomolets have shown that releases from the reactor have occurred and that the submarine has suffered considerable damage to its hulls. Norwegian monitoring detected 134Cs in surface sediments around Komsomolets in 1993 and 1994 and elevated activity concentrations of 137Cs in bottom seawater between 1991 and 1993. Since then and up to 2015, no increased activity concentrations of radionuclides above values typical for the Norwegian Sea have been observed in any environmental sample collected by Norwegian monitoring. In 2013 and 2015, Norwegian monitoring was carried out using an acoustic transponder on the sampling gear that allowed samples to be collected at precise locations, ∼20 m from the hull of Komsomolets. The observed 238Pu/239,240Pu activity ratios and 240Pu/239Pu atom ratios in surface sediments sampled close to Komsomolets in 2013 did not indicate any releases of Pu isotopes from reactor or the torpedo warheads. Rather, these values probably reflect the overprinting of global fallout ratios with fluxes of these Pu isotopes from long-range transport of authorised discharges from nuclear reprocessing facilities in Northern Europe. However, due to the depth at which Komsomolets lies, the collection of seawater and sediment samples in the immediate area around the submarine using traditional sampling techniques from surface vessels is not possible, even with the use of acoustic transponders. Further monitoring is required in order to have a clear understanding of the current status of Komsomolets as a potential source of radioactive contamination to the Norwegian marine environment. Such monitoring should involve the use of ROVs or submersibles in order to obtain samples next to and within the different compartments of the submarine.

Harvest-induced phenotypic selection in an island population of moose, Alces alces.

Empirical evidence strongly indicates that human exploitation has frequently led to rapid evolutionary changes in wild populations, yet the mechanisms involved are often poorly understood. Here, we applied a recently developed demographic framework for analyzing selection to data from a 20-year study of a wild population of moose, Alces alces. In this population, a genetic pedigree has been established all the way back to founders. We demonstrate harvest-induced directional selection for delayed birth dates in males and reduced body mass as calf in females. During the study period, birth date was delayed by 0.81 days per year for both sexes, whereas no significant changes occurred in calf body mass. Quantitative genetic analyses indicated that both traits harbored significant additive genetic variance. These results show that selective harvesting can induce strong selection that oppose natural selection. This may cause evolution of less favorable phenotypes that become maladaptive once harvesting ceases.

Outdoor thoron and progeny in a thorium rich area with old decommissioned mines and waste rock.

Radon (222Rn), thoron (220Rn) and their decay products may reach high levels in areas of high natural background radiation, with increased risk associated with mining areas. Historically, the focus has mostly been placed upon radon and progeny (RnP), but recently there have been reports of significant contributions to dose from thoron progeny (TnP). However, few direct measurements of TnP exist under outdoor conditions. Therefore, we assessed the outdoor activity concentrations of radon, thoron and TnP in an area of igneous bedrock with extreme levels of radionuclides in the thorium decay series. The area is characterized by decommissioned mines and waste rock deposits, which provide a large surface area for radon and thoron emanation and high porosity enhancing exhalation. Extreme levels of thorium and thoron have previously been reported from this area and to improve dose rate estimates we also measured TnP using filter sampling and time-integrating alpha track detectors. We found high to extreme levels of thoron and TnP and the associated dose rates relevant for inhalation were up to 8 µSvh-1 at 100 cm height. Taking gamma irradiation and RnP into account, significant combined doses may result from occupancies in this area. This applies to recreational use of the area and especially previous and planned road-works, which in the worst case could involve doses as large as 23.4 mSv y-1. However, radon and thoron levels were much more intense on a hot September day than during time-integrated measurements made the subsequent colder and wetter month, especially along the ground. This may be explained by cold air observed flowing out from inside the mines through a drainage pipe adjacent to the measurement stations. During warm periods, activity concentrations may therefore be due to both local exhalation from the ground and air ventilating from the mines. However, a substantially lower level of TnP was measured on the September day using filter sampling, as compared to what was measured with time-integrative alpha track detectors. A possible explanation could be reduced filter efficiency related to the attached progeny of some aerosol sizes, but a more likely cause is an upwards bias on TnP detectors associated with assumed deposition velocity, which may be different in outdoor conditions with wind or a larger fraction of unattached progeny. There is thus a need for better instrumentation when dealing with outdoor TnP.

Effects of inbreeding on fitness-related traits in a small isolated moose population.

Inbreeding can affect fitness-related traits at different life history stages and may interact with environmental variation to induce even larger effects. We used genetic parentage assignment based on 22 microsatellite loci to determine a 25 year long pedigree for a newly established island population of moose with 20-40 reproducing individuals annually. We used the pedigree to calculate individual inbreeding coefficients and examined for effects of individual inbreeding (f) and heterozygosity on fitness-related traits. We found negative effects of f on birth date, calf body mass and twinning rate. The relationship between f and calf body mass and twinning rate were found to be separate but weaker after accounting for birth date. We found no support for an inbreeding effect on the age-specific lifetime reproductive success of females. The influence of f on birth date was related to climatic conditions during the spring prior to birth, indicating that calves with a low f were born earlier after a cold spring than calves with high f. In years with a warm spring, calf f did not affect birth date. The results suggest that severe inbreeding in moose has both indirect effects on fitness through delayed birth and lower juvenile body mass, as well as separate direct effects, as there still was a significant relationship between f and twinning rate after accounting for birth date and body mass as calf. Consequently, severe inbreeding as found in the study population may have consequences for population growth and extinction risk.