Harbor seals, as generalist predators, can adjust their diet in response to shifts in prey availability; thus, their diet composition has the potential to act as a biodiversity indicator. This study examined the diet of harbor seals in the Skagerrak by analyzing otoliths from scat samples collected in 2020 and compared species composition to historical datasets from 1977/78 and 1989. The diet in 2020 encompassed 28 fish species. Sandeel and Norway pout were the most common prey, comprising 39.6% and 21.8% of otoliths, respectively. The group haddock/pollack/saithe and ballan wrasse had the highest relative biomass contributions: 18.7% and 13.5%, respectively. A comparison with historical data revealed a shift in diet, with a notable decline in cod and herring consumption and an increase in sandeel and haddock/pollack/saithe. The findings suggest a dietary adaptation of harbor seals to available prey species. The reduced intake of energy-rich prey like herring raises concerns about whether current diet composition can meet the seals' energetic needs. The observed dietary changes may reflect changes in prey availability due to environmental and anthropogenic factors, emphasizing the importance of continued studies to assess the implications for harbor seal populations in the region.

We present evidence that the abundance of harbour seals in the Kattegat-Skagerrak is in decline. Until recently, the Kattegat-Skagerrak harbour seal population has grown exponentially as it has recovered from historic over-hunting and two mass mortality events. This has provided an important case study for the influence of environmental factors on population growth. Over recent years, deteriorating average body condition and reduced pup counts in certain colonies have indicated that the population is under stress. At the same time, there has been an increase in hunting in the region which may compound existing environmental stressors. To determine trends in harbour seal abundance, we compile aerial survey data for the Kattegat-Skagerrak and South-Western Baltic Sea (S.W. Baltic) populations between the years 2003 and 2023. With parametric modelling, we find that a logistic growth model is the best fit for aerial survey data from the Kattegat-Skagerrak population, while an exponential growth model is the best fit for data from the smaller S.W. Baltic population. We determine trends by fitting non-parametric Generalised Additive Models (GAMs) to aerial survey data and calculating their first derivative. In the Kattegat-Skagerrak, we estimate an annual decline of - 408 individuals (SE = 242, CI95% = [- 882, 67]) at the end of the survey period. This decline represents approximately 3.3% of the estimated mean count in 2023. Rate of change remained positive in the separate S.W. Baltic population throughout the survey period at an estimated annual increase of 54 individuals (SE = 23, CI95% = [9,99]), representing approximately 3.9% of mean counts in 2023. We find that the final rate of change was zero or below zero in eleven out of twelve subregions (representing individual colonies) in the Kattegat-Skagerrak. Declining counts in the Kattegat-Skagerrak since the mid-2010s may be influenced by changes in haul-out behaviour as a result of increased population density or vessel traffic. Despite this, declines likely reflect real changes in seal abundance. Changes in abundance may be the result of well documented environmental degradation affecting prey availability and causing regionally lowered birth rates.

Extensive primary boreal forests within Europe are primarily located in Fennoscandia and northwestern Russia. These forests host numerous endemic and red-listed species but are rapidly being exploited and transformed to production forests that lack the habitat characteristics that are required for sustained biodiversity. Over the past 30 years, certain highly valuable areas within Swedish forests have been designated and registered as "woodland key habitats" (WKH) to be safeguarded from clear-cutting. However, despite their high conservation values, WKH lack proper legislative protection. Recently, many WKH were deregistered and thereby lost their potential protection against clear-cutting, jeopardizing biodiversity values in these forests. Moreover, the former way of classifying WKH has been criticized for being too lenient, making conventional forestry difficult. To assess the leniency of WKH registration and effect of WKH deregistration, we conducted a field inventory of WKH following a new inventory method proposed by the Swedish Forest Agency, featuring more stringent criteria for classification of WKH in north-western Sweden. The inventory was conducted in 9 still registered and 9 recently deregistered WKH to assess their conservation values. Our inventory results show that all 18 areas reach the criteria for WKH with the new method, despite higher, more stringent thresholds for conservation values. Hence, formerly registered WKH were not deregistered due to lower values. Moreover, analysis of recent harvest actions within deregistered WKH in Sweden showed that almost 1,200 hectares (~ 2%) of these areas were clear-cut or reported for clear-cutting 1-7 years post deregistration. As such, our results indicate that WKH contain high values, even using more stringent classification criteria, but also that deregistration of WKH does not consider these values and increases the risk of losing them to forest management. Given past and current declines in forest biodiversity, this is concerning, as conservation of areas containing high conservation values are needed, in order to preserve biodiversity in boreal forests.

The Baltic Sea is home to a genetically isolated and morphologically distinct grey seal population. This population has been the subject of 120-years of careful documentation, from detailed records of bounty statistics to annual monitoring of health and abundance. It has also been exposed to a range of well-documented stressors, including hunting, pollution and climate change. To investigate the vulnerability of marine mammal populations to multiple stressors, data series relating to the Baltic grey seal population size, hunt and health were compiled, vital demographic rates were estimated, and a detailed population model was constructed. The Baltic grey seal population fell from approximately 90,000 to as few as 3000 individuals during the 1900s as the result of hunting and pollution. Subsequently, the population has recovered to approximately 55,000 individuals. Fertility levels for mature females have increased from 9% in the 1970s to 86% at present. The recovery of the population has led to demands for increased hunting, resulting in a sudden increase in annual quotas from a few hundred to 3550 in 2020. Simultaneously, environmental changes, such as warmer winters and reduced prey availability due to overfishing, are likely impacting fecundity and health. Future population development is projected for a range of hunting and environmental stress scenarios, illustrating how hunting, in combination with environmental degradation, can lead to population collapse. The current combined hunting quotas of all Baltic Nations caused a 10% population decline within three generations in 100% of simulations. To enable continued recovery of the population, combined annual quotas of less than 1900 are needed, although this quota should be re-evaluated annually as monitoring of population size and seal health continues. Sustainable management of long-lived slowly growing species requires an understanding of the drivers of population growth and the repercussions of management decisions over many decades. The case of the Baltic grey seal illustrates how long-term ecological time series are pivotal in establishing historical baselines in population abundance and demography to inform sustainable management. Due to its isolation, the Baltic grey seal is subject to increased environmental stress relative to populations elsewhere. In this study, 120-years of data on this unique population are compiled and its future growth is modelled. Current hunting quotas in the Baltic are found to be unsustainable.image

Abstract Body mass is a fundamental indicator of animal health closely linked to survival and reproductive success. Systematic assessment of body mass for a large proportion of a population can allow early detection of changes likely to impact population growth, facilitating responsive management and a mechanistic understanding of ecological trends. One challenge with integrating body mass assessment into monitoring is sampling enough animals to detect trends and account for individual variation. Harbour seals (Phoca vitulina) are philopatric marine mammals responsive to regional environmental changes, resulting in their use as an indicator species. We present a novel method for the non‐invasive and semi‐automatic assessment of harbour seal body condition, using unoccupied aerial vehicles (UAVs/drones). Morphological parameters are automatically measured in georeferenced images and used to estimate volume, which is then translated to estimated mass. Remote observations of known individuals are utilized to calibrate the method. We achieve a high level of accuracy (mean absolute error of 4.5 kg or 10.5% for all seals and 3.2 kg or 12.7% for pups‐of‐the‐year). We systematically apply the method to wild seals during the Spring pupping season and Autumn over 2 years, achieving a near‐population‐level assessment for pups on land (82.5% measured). With reference to previous mark‐recapture work linking Autumn pup weights to survival, we estimate mean expected probability of over‐winter survival (mean = 0.89, standard deviation = 0.08). This work marks a significant step forward for the non‐invasive assessment of body condition in pinnipeds and could provide daily estimates of body mass for thousands of individuals. It can act as an early warning for deteriorating environmental conditions and be utilized as an integrative tool for wildlife monitoring. It also enables estimation of yearly variation in demographic rates which can be utilized in parameterizing models of population growth with relevance for conservation and evolutionary biology.

Changes in species abundance and distribution are driven by short-term interactions between individuals and their environment, as well as by long-term shifts in climate, prey abundance, disease outbreaks and predators, acting across generations. Unfortunately, management of wildlife populations by humans has often overlooked these long-term effects. This approach has resulted in an overestimation of nature’s resilience, leading to overharvesting and contributing to the decline and extinction of species. Field studies that document species demography over sufficiently long periods to understand the drivers of change are rare but crucial for sustainable management. In our view, these datasets should be recognised as World Heritage Ecological Time Series (WHETS). To counteract biodiversity loss, it is essential to document, preserve, and develop existing WHETS. We illustrate the value of such time series using the example of the Kattegat-Skagerrak Sea Region harbour seal (Phoca vitulina).

AbstractThe Centre for Marine Evolutionary Biology (CeMEB) at the University of Gothenburg, Sweden, was established in 2008 through a 10‐year research grant of 8.7 m€ to a team of senior researchers. Today, CeMEB members have contributed >500 scientific publications, 30 PhD theses and have organised 75 meetings and courses, including 18 three‐day meetings and four conferences. What are the footprints of CeMEB, and how will the centre continue to play a national and international role as an important node of marine evolutionary research? In this perspective article, we first look back over the 10 years of CeMEB activities and briefly survey some of the many achievements of CeMEB. We furthermore compare the initial goals, as formulated in the grant application, with what has been achieved, and discuss challenges and milestones along the way. Finally, we bring forward some general lessons that can be learnt from a research funding of this type, and we also look ahead, discussing how CeMEB’s achievements and lessons can be used as a springboard to the future of marine evolutionary biology.

Aerial line transect surveys of the density of humpback whales (Megaptera novaeangliae) conducted off West Greenland eight times between 1984 and 2007 were used to estimate the rate of increase on the summer feeding ground. Only surveys in 1993, 2005 and 2007 had enough sightings to construct independent density estimates, whereas the surveys in 1984–85 and 1987–89 had to be merged and treated as two surveys. The annual rate of increase was 9.4% yr–1 (SE = 0.01) between 1984 and 2007. This rate of increase is higher than the increase estimated at the breeding grounds in the West Indies, but is of the same magnitude as the observed rate of increase at other feeding grounds in the North Atlantic. A matrix model based on observed life history parameters revealed that the theoretical growth rate of a humpback whale population ranged between 1 and 11%. This confirms that the observed growth in West Greenland is within the plausible values. The survey in 2007 was used to make a fully corrected abundance estimate including corrections for whales that were submerged during the passage of the survey plane. The line transect estimate for 2007 was 1,020 (CV = 0.35). When the estimate was corrected for perception bias with mark-recapture distance sampling (MRDS) methods, the abundance increased to 1,505 (0.49). A correction for availability bias was developed based on time-depth-recorder information on the time spent at the surface (0–4m). However, used directly this correction leads to a positively-biased abundance estimate and instead a correction was developed for the non-instantaneous visual sighting process in an aircraft. The resulting estimate for 2007 was 3,272 (CV = 0.50) for the MRDS analysis. An alternative strip census estimate deploying a strip width of 300m resulted in 995 (0.33) whales. Correction for perception bias resulted in 991 (0.35) whales and corrected for the same availability bias as for the MRDS method resulted in a fully corrected estimate of 2,154 (0.36) humpback whales in West Greenland in 2007.

The harbour seal (Phoca vitulina) is the most widely distributed pinniped, occupying a wide variety of habitats and climatic zones across the Northern Hemisphere. Intriguingly, the harbour seal is also one of the most philopatric seals, raising questions as to how it colonized its current range. To shed light on the origin, remarkable range expansion, population structure and genetic diversity of this species, we used genotyping-by-sequencing to analyse ~13,500 biallelic single nucleotide polymorphisms from 286 individuals sampled from 22 localities across the species' range. Our results point to a Northeast Pacific origin of the harbour seal, colonization of the North Atlantic via the Canadian Arctic, and subsequent stepping-stone range expansions across the North Atlantic from North America to Europe, accompanied by a successive loss of genetic diversity. Our analyses further revealed a deep divergence between modern North Pacific and North Atlantic harbour seals, with finer-scale genetic structure at regional and local scales consistent with strong philopatry. The study provides new insights into the harbour seal's remarkable ability to colonize and adapt to a wide range of habitats. Furthermore, it has implications for current harbour seal subspecies delineations and highlights the need for international and national red lists and management plans to ensure the protection of genetically and demographically isolated populations.

Detecting changes in population trends depends on the accuracy of estimated mean population growth rates and thus the quality of input data. However, monitoring wildlife populations poses economic and logistic challenges especially in complex and remote habitats. Declines in wildlife populations can remain undetected for years unless effective monitoring techniques are developed, guiding appropriate management actions. We developed an automated survey workflow using unmanned aerial vehicles (drones) to quantify the number and size of individual animals, using the well-studied Scandinavian harbour seal (Phoca vitulina) as a model species. We compared ground-based counts using telescopes with manual flights, using a zoom photo/video, and pre-programmed flights producing orthomosaic photo maps. We used machine learning to identify and count both pups and older seals and we present a new method for measuring body size automatically. We evaluate the population’s reproductive success using drone data, historical counts and predictions from a Leslie matrix population model. The most accurate and time-efficient results were achieved by performing pre-programmed flights where individual seals are identified by machine learning and their body sizes are measured automatically. The accuracy of the machine learning detector was 95–97% and the classification error was 4.6 ± 2.9 for pups and 3.1 ± 2.1 for older seals during good light conditions. There was a clear distinction between the body sizes of pups and older seals during breeding time. We estimated 320 pups in the breeding season 2021 with the drone, which is well beyond the expected number, based on historical data on pup production. The new high quality data from the drone survey confirms earlier indications of a deteriorating reproductive rate in this important harbour seal colony. We show that aerial drones and machine learning are powerful tools for monitoring wildlife in inaccessible areas which can be used to assess annual recruitment and seasonal variations in body condition.

Empirical evidence has shown that historical exposure of polychlorinated biphenyls (PCBs) to Baltic grey seals not only severely affected individual fitness, but also population growth rates and most likely caused the retarded recovery rate of the depleted population for decades. We construct a new model which we term a toxicokinetic-toxicodynamic (TKTD) population model to quantify these effects. The toxicokinetic sub model describes in detail the bioaccumulation, elimination, and vertical transfer from mother to offspring of PCBs, and was linked to a toxicodynamic model for estimation of PCB-related damage, hazard, and stress impacts on fertility and survival rates. Both sub-models were then linked to a Leslie matrix population model to calculate changes in population growth rate and age structure given different rates of PCB exposure. Toxicodynamic model parameters related to reproductive organ lesions were calibrated using published historical data on observed pregnancy rates in Baltic grey seal females. Our model showed that increased PCB exposure caused reduced fertility, decreased vertical transfer, and increased biomagnification. Compared to empirical data, the TKTD population model described well the age-specific bioaccumulation pattern of PCBs in Baltic grey seals, and thus, the toxicokinetic parameters, deduced from literature, are believed to be reliable. The model also captured well the general effects of PCBs on historical population growth rates. The developed model can be used to perform population viability analyses of Baltic grey seals with multiple stressors, also including by-catches and different hunting regimes. The model can also be extended to other marine mammals and other contaminants than PCB by adjustments of model parameter values and thus provides a test bed in silico for new substances.

Generation time is a measure of the pace of life and is used to describe processes in population dynamics and evolution. We show that three commonly used mathematical definitions of generation time in age-structured populations can produce different estimates of up to several years for the same set of life history data. We present and prove a mathematical theorem that reveals a general order relation among the definitions. Furthermore, the exact population growth rate at the time of sampling influences estimates of generation time, which calls for attention. For phylogenetic estimates of divergence times between species, included demographic data should be collected when the population growth rate for each species is most common and typical. In conservation biology, demographic data should be collected during phases of population decline in declining species, contrary to common recommendations to use predisturbance data. The results can be used to improve the International Union for Conservation of Nature’s recommendation in parameterizing models for evaluating threat categories of threatened species and to avoid underestimating extinction risk.

This introductory chapter to our Environment International VSI does not need an abstract and therefore we just include our recommendations below in order to proceed with the resubmission. Future work should examine waterbirds as food web sentinels of multiple stressors as well as Baltic Sea food web dynamics of hazardous substances and how climate change may modify it. Also, future work should aim at further extending the new frameworks developed within BALTHEALTH for energy and contaminant transfer at the population level (Desforges et al., 2018, Cervin et al., 2020/this issue Silva et al., 2020/this issue) and their long term effects on Baltic Sea top predators, such as harbour porpoises, grey seals ringed seals, and white-tailed eagles. Likewise, the risk evaluation conducted for PCB in connection with mercury on Arctic wildlife (Dietz et al., 2019, not a BONUS BALTHEALTH product) could be planned for Baltic Sea molluscs, fish, bird and marine mammals in the future. Finally, future efforts could include stressors not covered by the BONUS BALTHEALTH project, such as food web fluxes, overexploitation, bycatches, eutrophication and underwater noise.

Graphical AbstractPOPs in Antarctic Mammals.

Conservation efforts have mainly been focused on depleted species or populations, but many formerly reduced marine mammal populations have recovered to historical abundances. This calls for new management strategies and new models for ecological risk assessment that incorporate local density dependence and multiple environmental stressors. The harbor seal metapopulation in Swedish and Danish waters has increased from about 2500 to 25,000 over the past 40 yr. Trend analysis based on aerial survey data and somatic growth curves indicates that the population is close to carrying capacity. We performed a population viability analysis based on realistic life history parameters and investigated a range of potential scenarios caused by future stressors. If the population is able to resume its high intrinsic rate of increase at about 11% annually, when pushed down below carrying capacity, it can also sustain additional mortality such as modest hunting and infrequent epizootics. However, if xenobiotics will cause even a slight reduction in average fecundity, the population becomes significantly more vulnerable. In the absence of epizootics, and given full reproductive capacity, hunting of a few hundred animals annually is not harmful to the long-term persistence of the population. Nevertheless, a slight decrease in growth potential, for example, caused by exposure to endocrine disruptors, makes even limited hunting risky. Our study shows how an apparently stable and abundant marine mammal population can be close to a point of rapid population decline. Thus, careful monitoring of population size, growth rate, health, and exposure to xenobiotics as well as recording of the age and sex structure of the hunt is required to avoid repeating the history of overexploitation and another population collapse.

Wildlife population dynamics are shaped by multiple natural and anthropogenic factors, including predation, competition, stressful life history events, and external environmental stressors such as diseases and pollution. Marine mammals such as gray seals rely on extensive blubber layers for insulation and energy storage, making this tissue critical for survival and reproduction. This lipid rich blubber layer also accumulates hazardous fat soluble pollutants, such as polychlorinated biphenyls (PCBs), that can directly impact adipose function or be mobilized during periods of negative energy balance or transferred to offspring to exert further impacts on target tissues or vulnerable life stages. To predict how marine mammals will respond to ecological and anthropogenic stressors, it is necessary to use process-based modelling approaches that integrate environmental inputs, full species life history, and stressor impacts with individual dynamics of energy intake, storage, and utilization. The purpose of this study was to develop a full lifecycle dynamic energy budget and individual based model (DEB-IBM) that captured Baltic gray seal physiology and life history, and showcase potential applications of the model to predict population responses to select stressors known to threaten gray seals and other marine mammals around the world. We explore variations of three ecologically important stressors using phenomenological simulations: food limitation, endocrine disrupting chemicals that reduce fertility, and infectious disease. Using our calibrated DEB-IBM for Baltic gray seals, we found that continuous incremental food limitation can be more detrimental to population size than short random events of starvation, and further, that the effect of endocrine disruptors on population growth and structure is delayed due to bioaccumulation, and that communicable diseases significantly decrease population growth even when spillover events are relatively less frequent. One important finding is the delayed effect on population growth rate from some stressors, several years after the exposure period, resulting from a decline in somatic growth, increased age at maturation and decreased fecundity. Such delayed responses are ignored in current models of population viability and can be important in the correct assessment of population extinction risks. The model presented here provides a test bed on which effects of new hazardous substances and different scenarios of future environmental change affecting food availability and/or seal energetic demands can be investigated. Thus, the framework provides a tool for better understanding how diverse environmental stressors affect marine mammal populations and can be used to guide scientifically based management.

Many aspects of sexual and asexual reproduction have been studied empirically and theoretically. The differences between sexual and asexual reproduction within a species often lead to a biased geographical distribution of individuals with different reproductive strategies. While sexuals are more abundant in the core habitat, asexuals are often found in marginal habitats along the edge of the species distribution. This pattern, called geographic parthenogenesis, has been observed in many species but the mechanisms reponsible for generating it are poorly known. We used a quantitative approach using a metapopulation model to explore the ecological processes that can lead to geographic parthenogenesis and the invasion of new habitats by different reproductive strategies. We analyzed the Allee effect on sexual populations and the population sensitivity to environmental stress during the invasion of a marginal, unstable habitat to demonstrate that a complex interaction between the Allee effect, sensitivity to environmental stress and the environmental conditions can determine the relative success of competing reproductive strategies during the initial invasion and longterm establishment in the marginal habitat. We discuss our results in the light of previous empirical and theoretical studies. Author Summary Individuals can reproduce with or without sex. Very often, closely related species are distributed in a such a way that the sexually reproducing species is most frequently found in the core habitat while the asexually reproducing species is found on the edge of the habitat range. This biased distribution of reproductive strategies across a habitat range is called geographic parthenogenesis and has been observed in several species. While many processes have been proposed to explain such a pattern, a quantitative approach of the ecological processes was absent. We investigated important differences between sexual and asexual reproduction and how these differences affect the success of sexuals and asexuals invading a marginal, unstable environment. We showed that the relative frequency of each reproductive strategy in the marginal habitat depends on how much sexuals rely on population density to reproduce and how much asexuals are affected by environmental stress relative to sexuals. Our study presents a quantitative ecological explanation for geographic parthenogenesis and provides the conditions under which different distribution patterns can emerge.

Here we review contaminant exposure and related health effects in six selected Baltic key species. Sentinel species included are common eider, white-tailed eagle, harbour porpoise, harbour seal, ringed seal and grey seal. The review represents the first attempt of summarizing available information and baseline data for these bio-monitoring key species exposed to industrial hazardous substances focusing on anthropogenic persistent organic pollutants (POPs). There was only limited information available for white-tailed eagles and common eider while extensive information exist on POP exposure and health effects in the four marine mammal species. Here we report organ-tissue endpoints (pathologies) and multiple biomarkers used to evaluate health and exposure of key species to POPs, respectively, over the past several decades during which episodes of significant population declines have been reported. Our review shows that POP exposure affects the reproductive system and survival through immune suppression and endocrine disruption, which have led to population-level effects on seals and white-tailed eagles in the Baltic. It is notable that many legacy contaminants, which have been banned for decades, still appear to affect Baltic wildlife. With respect to common eiders, changes in food composition, quality and contaminant exposure seem to have population effects which need to be investigated further, especially during the incubation period where the birds fast. Since new industrial contaminants continuously leak into the environment, we recommend continued monitoring of them in sentinel species in the Baltic, identifying possible effects linked to climate change, and modelling of population level effects of contaminants and climate change.

Many endangered marine mammal populations are difficult to study, spread out over large areas, and capturing them for branding and research purposes would be unethical. Yet, they are in urgent need for assessment and conservation actions. We suggest collecting data from other more abundant populations of the same species, with careful consideration of body size, age at sexual maturity, and ecological conditions to produce scientifically sound best approximations of vital rates of data deficient endangered populations. The genetically distinct Baltic Sea harbour porpoise population amounts to about 500 animals and is classified as ‘Critically Endangered’ according to the IUCN red list. Data deficiency on nearly all demographic parameters have precluded systematic investigations of the relative importance of stressors affecting population viability. We took a comparative life history approach and investigated the phenotypic plasticity in somatic and demographic vital rates of seven larger, well studied North Atlantic harbour porpoise populations, enabling us to approximate the missing pieces of the life history of the Baltic population. We parameterized a stochastic, individual-based population model, and performed a population viability analysis for a range of biologically realistic parameter values and scenarios of environmental stressors. The baseline scenario was based on the most representative samples of healthy harbour porpoise populations and challenged with three different levels of bycatches. Due to high levels of endocrine disruptive contaminants observed in Baltic harbour porpoises, we also investigated the effect of a possible reduction in fecundity. Subsequently, the combined effects of bycatches and reduced fecundity were investigated in terms of population growth rate and quasi-extinction risk. The Baltic harbour porpoise population is viable in the baseline scenario without anthropogenic stressors. However, even the lowest estimated bycatch level of 7 individuals per year will lead to a population collapse to ≤50 animals with high probability (0.4–1.0) over the next century, assuming an intermediate or low (<73%) fecundity. Adult survival is of critical importance and mitigation of fishery impacts and reduction of anthropogenic disturbances in the identified main breeding areas are recommended.