Human land use can take advantage of using exotic species to increase financial benefits. However, the use of exotic tree species might affect ecosystem functioning, potentially including the habitat use and movement behaviour of animals, modifying their ecological impact, and interactions with human land use. The native pine of Sweden is Scots pine Pinus sylvestris. In 1920, the Swedish forestry introduced the North American lodgepole pine P. contorta to Sweden to increase timber production.We used a two-year data set from GPS-collared adult moose Alces alces to examine the annual and seasonal habitat selection of this large keystone herbivore in a managed forest landscape with a relatively higher percentage of P. contorta compared to other Swedish boreal regions. Compared to other studies, a central departure point in this work was our focus on whether moose exhibited different selection between the native P. sylvestris stands and the exotic P. contorta stands.We found no evidence that moose avoided stands of P. contorta within forest landscapes. Instead, our results show that young stands of both P. sylvestris and P. contorta were a highly preferred habitat for moose year-round, and that selection for P. contorta forests depended on the proportion of P. contorta at the landscape level and the season. Our work also reveals a diverse selection of forested habitats in almost all seasons, reflecting the differing needs of moose during different times of the year.Despite being rare in our system, we suggest our findings can have important implications for forestry - P. contorta is often planted in the belief that these forests less likely to be browsed by moose than are native P. sylvestris forests. Our results do not indicate that the current distribution of different P. contorta stands has a limiting effect on how moose distribute across the landscape matrix.

Ungulate browsing is widely regarded as a major constraint on the recruitment of broadleaves important as carriers of biodiversity. Here, we combine observational data from 123 Moose Management Areas (MMAs) and 30 experimental exclosure pairs to evaluate the recruitment targets set by the Swedish Forest Agency and Swedish Environmental Protection Agency for rowan (Sorbus aucuparia), aspen (Populus tremula), willows (Salix spp.), and oaks (Quercus spp.) (RAWO) in Swedish moose management. We investigated goal fulfilment and predictors of recruitment using national monitoring data. Targets were met for only similar to 20% of Sweden. Site productivity (+) was the strongest predictor of recruitment, while moose and deer densities had little influence. Experimental exclusions of browsers primarily affected the transition of seedlings into competitive size classes, not initial establishment, with growth stalling after the first 3-4 years when browsers were present. In an experiment on intermediate-productivity soils, RAWO grew slower than pine and recruitment fell below targets, even in exclosures without ungulate browsing. In a second experiment on high-productivity soils, browsing constrained RAWO growth but competitive RAWO stem density remained well above the target, even with browsing. Without browsing, combined measures of RAWO kept up with conifers. Responses to browsing and competition differed between RAWO species, such that rowan, willow and oak were more affected than aspen. Our results indicate that while browsing limits RAWO recruitment, site productivity and competition are also important. We suggest that the current quantitative targets may have little relevance and may be inappropriate for setting harvest quotas in MMAs in Sweden.

Reproductive success in seasonal environments depends on aligning birth and early offspring development with favourable ecological conditions. In mammals with embryonic diapause, birth timing emerges from a sequence that includes reproductive activation, mating and ovulation, diapause termination, implantation and parturition. Embryonic diapause can allow mating and birth to occur under different favourable conditions, but whether birth timing adjusts to directional environmental change depends on the plasticity of the reproductive stages between mating and parturition. We use mustelids as a comparative system to develop a stage-specific framework to identify when diapause buffers environmental variation and when it limits adjustment to shifting ecological optima. The framework distinguishes experimentally supported mechanisms and observed reproductive patterns from climate-related hypotheses that remain to be tested. Photoperiod provides a predictable seasonal signal, whereas temperature-sensitive ecological conditions may change independently of daylength. Mustelids offer comparative leverage because closely related species differ in the presence and duration of diapause, ovulatory mechanisms, and regulation of reproductive timing. We identify three testable hypotheses: (1) responsiveness of birth timing depends on the reproductive stage at which environmental or physiological information modifies the mating-to-birth interval; (2) phenological mismatch arises when realised birth timing diverges from the timing associated with highest reproductive success; and (3) diapause influences both the transmission of variation from mating to birth and the capacity of birth timing to track directional environmental change. Evidence that climate-sensitive ecological changes shift the optimal timing of birth and reduce reproductive success when births fail to track that shift remains limited in mustelids. Testing these hypotheses requires reproductive-stage timing, ecological conditions and reproductive outcomes to be measured within the same populations.

Natural disturbances, including herbivory by deer, are key drivers of forest dynamics, yet their role in shaping field-layer plant diversity remains unresolved. We investigated how variation in deer density and community composition relates to field-layer plant richness in boreonemoral forest, with reference to the Intermediate Disturbance Hypothesis (IDH). Using annual pellet counts from 2012 to 2023, we quantified long-term densities of moose (Alces alces), red deer (Cervus elaphus), and small deer (roe Capreolus capreolus and fallow Dama dama combined) across 33 1 x 1 km forest tracts in central Sweden. We derived a Deer Energetic Requirement (DER) index by converting species densities to Klieber-scaled metabolic demands and summing across species. In 2023, field-layer vascular plants were surveyed, and relationships between deer densities, vegetation structure, and species richness were analysed. Bilberry browsing increased with both small deer density and DER, confirming that density estimates reflect realised browsing pressure. Vascular plant richness showed a unimodal relationship with small deer density, consistent with the IDH. Contrastingly, field-layer richness had a negative relationship with moose population density, while no relationship was found for red deer. Intermediate levels of DER were associated with lower dwarf shrub cover and higher vertical gap fraction and graminoid cover. These structural differences appeared to be the main indirect pathway through which deer density was related to diversity. Our findings emphasise the importance of browser community composition, density, and heterogeneity in structuring boreonemoral field-layers. Moderate browsing, especially by small deer, can enhance plant diversity by mediating resource availability in forest ecosystems.

Efforts to enhance forest biodiversity, climate resilience, and multifunctionality have renewed interest in harvesting methods that complement or replace clear-felling. In countries such as Sweden, where clear-felling remains dominant, understanding the behavioural mechanisms shaping forest owners' harvesting intentions is essential for guiding policy change. This study applies the Theory of Planned Behaviour (TPB) to examine the relative importance of attitudes, subjective norms, and perceived behavioural control in explaining Swedish private forest owners' intentions to apply clear-felling and alternative harvesting methods. Using survey data from 3095 respondents and structural equation modelling (SEM), we find that attitudes and perceived behavioural control are the strongest predictors of harvesting intentions, while subjective norms were not significant predictors in the models. Together, these factors explain a large share of the variance in harvesting intentions (R2 = 0.51–0.62). Complementary multiple linear regression analyses show that past experiences, professional advice, and economic considerations substantially influence attitudes and perceived behavioural control, acting as both barriers and enablers of change. Although clear-felling remains economically and structurally favoured, forest owners report similar levels of intention to apply alternative harvesting methods, indicating an openness to diversifying harvesting practices in Sweden. Targeted policy interventions that combine financial incentives with advisory and peer-learning support are likely needed to translate these intentions into practice. The results highlight that the foundations for a more diversified and sustainable forestry are already emerging among private forest owners, if policy and advisory systems can effectively nurture and sustain this momentum.

Drönare utrustade med termiska kameror har under senare år utvecklats till ett potentiellt verktyg för övervakning av klövvilt. Syftet med denna studie var att genomföra en första utvärdering av drönarbaserad inventering av älg för att på sikt kunna utveckla rekommendationer för en kvalitetssäkrad metodik inom svensk viltförvaltning. Studien genomfördes på uppdrag av Naturvårdsverket och omfattade inventeringar i tre studieområden i norra och mellersta Sverige. Resultaten visade att både manuella och automatiserade inventeringar med drönare kan användas för att uppskatta älgtäthet. Under de kalla och molniga vinterförhållandena som rådde vid inventeringarna skattades upptäckningsgraden till nära 100 %. Inventeringar av GPS-märkta älgar och parallella flygningar genomförda av olika piloter visade att i princip samtliga älgar som fanns inom inventeringsområdena kunde återfinnas. Simuleringar baserade på totalinventeringar visade att stickprovsbaserade inventeringar med provytor kan ge användbara täthetsskattningar, men att den nödvändiga täckningsgraden varierar beroende på hur älgarna är fördelade i landskapet. I områden där älgarna var måttligt koncentrerade i landskapet bedöms cirka 20–30 % av arealen ofta vara tillräckligt, medan upp till 50 % kan behöva inventeras i områden där älgarna uppträder mer koncentrerat. För skattning av könskvot och antal kalvar per hondjur krävs generellt större stickprov än för att enbart skatta täthet. Studien visar också att det är möjligt att bestämma kön och ålder hos observerade älgar med hjälp av zoombilder, men att detta kräver mer tid, högkvalitativ bildinsamling och erfaren bedömning. Viktiga begränsningar utgörs av väderberoende, drönarnas räckvidd och nuvarande regelverk för flygning inom synhåll (VLOS). För att möjliggöra effektiv inventering av större områden rekommenderas fortsatt utveckling av metoder för flygning bortom visuell siktlinje (BVLOS), liksom standardisering av metodik, utbildning och kvalitetssäkring. Sammantaget visar resultaten att drönare har stor potential som ett komplement till befintliga metoder för älgövervakning och viltförvaltning.

Human–wildlife interactions present both challenges and opportunities across Europe. Managing these interactions often requires decisions that fall between reactive (taking action after a problem arises) and proactive (anticipating and preventing negative impacts) management. This study examines factors influencing public attitudes toward proactive and reactive wildlife management in six European countries (France, Germany, Italy, Spain, Sweden and United Kingdom), based on an online survey of 2400 participants. We relied on the cognitive hierarchy framework to explain relationships between management attitudes and factors such as wildlife value orientations, risk perceptions, and perceptions of red deer (Cervus elaphus L.). Attitudes toward wildlife management were generally shaped by value orientations and risk perception. Mutualism was associated with support for proactive wildlife and human protection, whereas domination was associated with reactive and proactive management aimed at protecting human interests. Higher perceived risk further increased support for proactive management strategies. These results provide practical insights into how public attitudes relate to wildlife management.

Across the northern hemisphere, ungulates are expanding in range and abundance, forming novel communities in increasingly human-modified landscapes. These shifts drive new interactions over available food resources, but patterns of resource use and partitioning in Europe's multi-species systems remain poorly understood. This study examined seasonal diets and resource partitioning in diverse cervid communities (moose, roe deer, red deer, and fallow deer) across two Swedish landscapes (coastal-boreal and boreo-nemoral) differing in deer density and land use. Based on their foraging strategies, we expected (Hypothesis 1) diet richness and dietary niche width to be greater in intermediate feeders (red and fallow deer) than in browsers (moose and roe deer), (Hypothesis 2) trophic partitioning between browsers and intermediate feeders to be driven mainly by graminoid use, and (Hypothesis 3) intra- and interspecific overlap to vary with season, deer density, habitat diversity, and proportion of arable land. DNA metabarcoding of 2568 fecal samples showed that deer consumed plants from over 70 families, though diets were typically dominated by fewer than 10. Vaccinium shrubs were key forages year-round, while birch and willow dominated during the growing season. Moose consumed large amounts of pine in spring and winter (> 50% in the boreo-nemoral, 35%-40% in the coastal-boreal landscape), with less during summer-autumn (~15%). Forbs were important for smaller deer, especially in spring and summer-autumn, and more heavily used in winter in the boreo-nemoral landscape, likely due to supplementary feeding with human-provided food like hay or silage. Spruce use was low overall (< 5%), with fallow deer showing the highest intake. Consistent with Hypothesis 1, diet richness and niche width increased from moose to fallow deer. In partial support of Hypothesis 2, principal coordinates analysis (PCoA) revealed that graminoids contributed to trophic partitioning, but the pattern was not a strict browser-intermediate feeder divide. Moose consistently separated from the smaller deer due to avoidance of graminoids and reliance on pine and juniper, while roe deer, although a browser, sometimes overlapped with red and fallow deer through greater use of graminoids. During winter in the coastal-boreal landscape, wavy hairgrass (Avenella flexuosa) contributed to the significant separation between browsing roe deer and intermediate-feeding red deer diets, consistent with Hypothesis 2. Diet overlap among smaller deer varied with season and landscape. Intraspecific overlap was the highest in moose and the lowest in fallow deer, declining during summer-autumn across species. Overlap was influenced by deer density, habitat diversity, and arable land, consistent with Hypothesis 3, but effects were species-specific and explained only limited variation. Our results highlight the dietary plasticity of red and fallow deer, which may intensify resource competition with moose and roe deer in multi-species systems, particularly where supplementary feeding is common. These insights support adaptive, multi-species management of deer in northern ecosystems.

Knowledge about habitat selection is crucial for ungulate management and conservation to handle competing land use. Thermal tolerance and access to food are two important drivers of habitat selection. Thus, moving into thermal shelter when temperatures increase may result in reduced energy intake e.g. mature forests providing thermal shelters through high canopy cover, but lower amounts of forage than more open habitats. Body size affects heat sensitivity, with larger animals being more sensitive. In this study, we investigated ambient temperature mediated habitat selection in three differently-sized, sympatric, deer species (moose, red deer and roe deer) during spring and summer in Sweden. We also assessed the trade-off between thermal shelter and forage availability. We used data from GPS-collared animals with a combination of land cover and airborne LIDAR data to quantify canopy cover as a proxy for thermal refuge and shrub cover as a proxy for forage availability. We found that temperature influenced habitat selection in all three deer species however with divergent patterns. During warmer daily temperatures, moose and red deer selected areas with more canopy cover for thermal shelter, however showing opposite patterns during warmer nights, suggesting patterns of thermoregulatory behaviour. Roe deer showed no influence of high temperatures on canopy cover selection however,, selected more strongly for areas with high canopy cover during colder temperatures i.e. showing contrasting patterns compared to moose and red deer, suggesting that canopy cover may be relatively less important for roe deer as temperature increase. All three species selected for shrub cover during warmer temperatures, which can provide both high forage availability and shade over bed sites for smaller deer species. Our findings indicate that canopy cover and shrub cover habitats appear to be important as temperatures increase, but their significance varies among species. Furthermore, our study highlights the complexity behind habitat selection in sympatric ungulate species, food intake and predation risk.

Cervids inhabit mosaic landscapes that are strongly influenced by human land use and actions. Forestry and agriculture influence both the foodscape and the risk landscape, thus influencing how cervids respond to resource-risk trade-offs. Furthermore, these land use types interact to shape cervid use of the landscape. For example, cervids can forage in open agricultural fields providing high quality forage during the night, while using nearby forest patches to reduce risk during the day. Hence, actions conducted in one of these land use types will likely influence the use of the other by cervids. In this study, we investigate how agricultural and silvicultural practices interact to shape cervid use of the forest and their impact on commercially valuable trees. More specifically, we investigate the effect of growing nutrient dense crops in mixed forest-agriculture landscapes. We hypothesized that the sowing of such crops may not only attract cervids to the agricultural parts of the landscape, but also lead to increased browsing levels in adjacent forest areas where cervids seek cover. Moreover, such high quality crops may not only influence the number of animals, but also increase the intake rate of fibrous tree saplings of individual cervids to nutritionally balance the high input of easily-digestible macronutrients from the crops. We tested these ideas by conducting a large landscape-scale experiment where we manipulated the foodscape by growing two crop types on 28 fields (14 fields with high-quality oat versus 14 fields with lower-quality grass) and measuring cervid use of tree saplings in the forest patches adjacent these fields. We also investigated how forage availability in the forest patches and the distance from field edge affected browsing on tree saplings. Finally, we compared the effects of crop type and forest forage availability on the browsing pressure on coniferous versus deciduous tree species. We studied this in a multispecies ungulate system, where four cervid species occur sympatrically. We show that the type of crop sown influenced browsing pressure on tree saplings in nearby forests, with higher browsing levels in forests surrounding oat compared to grass fields. Moreover, browsing pressure in these forests was influenced by the availability of young trees in the forest stands, where more young trees led to reduced browsing pressure on individual saplings. Our study emphasizes that human practices that shape features of the foodscape in one land use type will influence the impact cervids have in another land use type. Hence, our study highlights the importance of considering the actions and impacts of different land uses simultaneously for predicting cervid foraging patterns in mixed forest-agriculture landscapes.

Abstract The term “applied ecology of fear” was recently introduced to describe the growing research field that applies the theory of the ecology of fear to manage wildlife behaviour. The management goal is to drive targeted species spatially and temporally away from areas of human interest by inducing cues from real or simulated predators to reduce human‐wildlife conflict. We aimed to quantify, through a meta‐analysis, if prey anti‐predator response would vary among field trials versus pen‐based studies, predator cue types, predator hunting style and prey feeding type, and be stronger in response to larger predators relative to the prey's size. We also explored what studies found in terms of wildlife habituation to cues. We used species belonging to the Cervidae family as a case study since deer are among the group of species with the highest degree of human‐wildlife conflict. We retrieved 114 studies from online databases and collected information from 39 of those studies that fitted our research scope. We found that acoustic cues more frequently led to an anti‐predator response in deer than olfactory or visual cues. Neither predator hunting strategy nor deer feeding strategy or type of study (free‐ranging or pen‐based animals) influenced the extent to which deer responded to cues. Deer more frequently responded to cues that belonged to a larger predator relative to their size. Habituation was reported in less than one‐third of the studies, with a study period ranging from 1 to 90 days, and occurred as soon as 7 days after the start of the study on average. Our meta‐analysis suggested that acoustic cues hold most potential as a tool to manage deer behaviour. These findings support the development of applied ecology of fear tools that introduce predator cues to reduce human‐wildlife conflicts. Major knowledge gaps remain that limit the effective use of such tools in wildlife management and future research should focus on improving our understanding of habituation to cues, on comparing the effectiveness of different types of cues, on simultaneously using a combination of cue types, and on testing cues at spatial–temporal scales of actual land‐uses.

A paradox in evolutionary biology is how supergenes can maintain high fitness despite reduced effective population size, the suppression of recombination, and the expected accumulation of mutational load. The ruff supergene involves 2 rare inversion haplotypes (satellite and faeder). These are recessive lethals but with dominant effects on male mating strategies, plumage, and body size. Sequence divergence to the wild-type (independent) haplotype indicates that the inversion could be as old as 4 million years. Here, we have constructed a highly contiguous genome assembly of the inversion region for both the independent and satellite haplotypes. Based on the new data, we estimate that the recombination event(s) creating the satellite haplotype occurred only about 70,000 yr ago. Contrary to expectations for supergenes, we find no substantial expansion of repeats and only a modest mutation load on the satellite and faeder haplotypes despite high sequence divergence to the non-inverted haplotype (1.46%). The essential centromere protein N (CENPN) gene is disrupted by the inversion and is as well conserved on the inversion haplotypes as on the noninversion haplotype. These results suggest that the inversion may be much younger than previously thought. The low mutation load, despite recessive lethality, may be explained by the introgression of the inversion from a now extinct lineage.

Foraging on crops by wild ungulates may create human–wildlife conflicts through reducing crop production. Ungulates interact with and within complex socio‐ecological systems, making the reduction of crop damage a challenging task. Aside from ungulate densities, crop damage is influenced by different drivers affecting ungulate foraging behaviour: food availability and food quality in the landscape (i.e. the foodscape) as well as fear from hunting and scaring actions (i.e. the landscape of fear) may together affect the degree of damage via both direct and indirect effects. A better understanding of the individual effects of these potential drivers behind crop damage is needed, as is an appreciation of whether the effects are dependent on ungulate density. We investigated this by applying path analysis to test indirect and direct links between ungulate density, foodscape, landscape of fear and human management goals on crop damage of oats and grass, respectively. Our results suggest that crop type is the major driver behind crop damage, with more damage to oats than to leys, implying that human decisions (i.e. changing crop type) influence the level of crop damage. We found that management goals and actions influenced the foodscape and the landscape of fear, by affecting the amount of forage produced in the agricultural landscape and the amount of scaring actions. Additionally, we found that supplementary feeding influenced the local ungulate densities in the area. Our results highlight the importance of including human actions on multiple levels when assessing drivers behind damage by ungulates in managed landscapes. We suggest that more studies using path analysis on multiple scales are needed in order to tackle complex issues, such as crop damage and other human–wildlife conflicts.

Climate change represents a growing ecological challenge. The (sub) arctic and boreal regions of the world experience the most rapid warming, presenting an excellent model system for studying how climate change affects mammals. Moose ( Alces alces ) are a particularly relevant model species with their circumpolar range. Population declines across the southern edge of this range are linked to rising temperatures. Using a long-term dataset (1988–1997, 2017–2019), we examine the relative strength of direct (thermoregulatory costs) and indirect (food quality) pathways linking temperature, precipitation, and the quality of two important food items (birch and fireweed) to variation in moose calf mass in northern Sweden. The direct effects of temperature consistently showed stronger relationships to moose calf mass than did the indirect effects. The proportion of growing season days where the temperature exceeded a 20 °C threshold showed stronger direct negative relationships to moose calf mass than did mean temperature values. Finally, while annual forb (fireweed) quality was more strongly influenced by temperature and precipitation than were perennial (birch) leaves, this did not translate into a stronger relationship to moose calf weight. The only indirect path with supporting evidence suggested that mean growing season temperatures were positively associated with neutral detergent fiber, which was, in turn, negatively associated with calf mass. While indirect impacts of climate change deserve further investigation, it is important to recognize the large direct impacts of temperature on cold-adapted species.

The moose Alces alces is the largest herbivore in the boreal forest biome, where it can have dramatic impacts on ecosystem structure and dynamics. Despite the importance of the boreal forest biome in global carbon cycling, the impacts of moose have only been studied in disparate regional exclosure experiments, leading to calls for common analyses across a biome-wide network of moose exclosures.In this study, we use airborne laser scanning (ALS) to analyse forest canopy responses to moose across 100 paired exclosure-control experimental plots distributed across the boreal biome, including sites in the United States (Isle Royale), Canada (Quebec, Newfoundland), Norway, Sweden and Finland.We test the hypotheses that canopy height, vertical complexity and above-ground biomass (AGB) are all reduced by moose and that the impacts vary with moose density, productivity, temperature and pulse disturbances such as logging and insect outbreaks.We find a surprising convergence in forest canopy response to moose. Moose had negative impacts on canopy height, complexity and AGB as expected. The responses of canopy complexity and AGB were consistent across regions and did not vary along environmental gradients. The difference in canopy height between exclosures and open plots was on average 6 cm per year since the start of exclosure treatment (+/- 2.1 SD). This rate increased with temperature, but only when moose density was high.The difference in AGB between moose exclosures and open plots was 0.306 Mg ha(-1) year(-1) (+/- 0.079). In browsed plots, stand AGB was 32% of that in the exclosures, a difference of 2.09 Mg ha(-1). The uniform response allows scaling of the estimate to a biome-wide impact of moose of the loss of 448 (+/- 115) Tg per year, or 224 Tg of carbon.Synthesis: Analysis of ALS data from distributed exclosure experiments identified a largely uniform response of forest canopies to moose across regions, facilitating scaling of moose impacts across the whole biome. This is an important step towards incorporating the effect of the largest boreal herbivore on the carbon cycling of one of the world's largest terrestrial biomes.

1. Differences in botanical diet compositions among a large number of moose faecal samples collected during winter correlated with the nutritional differences identified in the same samples (Mantel-r = 0.89, p = 0.001), but the nutritional differences were significantly smaller (p < 0.001). 2. Nutritional geometry revealed that moose mixed Scots pine Pinus sylvestris and Vaccinium spp. as nutritionally complementary foods to reach a nutritional target resembling Salix spp. twigs, and selected for Salix spp. browse (Jacob's D > 0). 3. Available protein (AP) and total non-structural carbohydrates (TNC) were significantly correlated in observed diets but not in hypothetical diets based on food availability. 4. The level of Acetoacetate in moose serum (i.e. 'starvation') was weakly negatively associated with digestibility of diets (p = 0.08) and unrelated to increasing AP:TNC and AP:NDF ratios in diets (p > 0.1). 5. Our study is the first to demonstrate complementary feeding in free-ranging moose to attain a nutritional target that has previously been suggested in a feeding trial with captive moose. Our results add support to the hypothesis of nutritional balancing as a driver in the nutritional strategy of moose with implications for both the management of moose and food resources.

Populations of large herbivores, including members of the deer family Cervidae, are expanding across and within many regions of the northern hemisphere. Because their browsing on trees can result in economic losses to forestry and strongly affect ecosystems, it is becoming increasingly important to understand how best to mitigate resultant damage. Previous research has highlighted the importance of regulating deer density and the availability of alternative forage to reduce browsing damage levels in conifer production stands. However, often only one or two proxies of forage availability have been used instead of applying a broad foodscape approach and more knowledge is needed to understand which types of alternative forage best mitigate damage. We conducted field inventories of damage that occurred during the previous fall/winter in 112 production stands in southern Sweden, while also measuring forage availability and cervid faecal pellets in the surrounding landscape (16 ha). Local landowners provided data on supplementary feeding. We found that variation in cervid (Alces alces, Capreolus capreolus, Cervus elaphus and Dama dama) browsing damage to top shoots or stems of young Scots pine trees (Pinus sylvestris, hereon pine), was better explained by the availability of alternative natural forage (using several indices and species of trees and shrubs) than by supplementary feeding. The proportion of damaged pine trees was higher in stands with a lower density of pine stems; in landscapes with a lower density of key broadleaf tree species (genera Sorbus, Salix, Populus and Quercus); and in landscapes with more open land (agricultural fields and paddocks). Damage was also higher in stands where relatively large amounts of moose faeces was found, while not related to the amount of faeces from other cervid species. The amount of supplementary feed (silage or other types such as root vegetables) did not explain variation in pine damage, but the result was possibly affected by relatively few study areas supplying sufficient data on supplementary feeding. The results from our inventory illustrate the efficacy of using naturally growing forage to mitigate browsing damage to young pine trees in managed landscapes. Creation of such forage is also recommended over supplementary feeding because of co-benefits to forest biodiversity and ecosystem services.

1.European large herbivore populationsincreased dramatically during the past decades. As a result, interactions among different large herbivore species, and land-use, such as forestry and agriculture, are also intensifying. However, there is little ecological knowledge on how the surrounding human-dominated landscape shapes interactions between these species as well as ecosystem services such as forestry, derived from the landscape. A better understanding of landscape scale mechanisms driving herbivore foraging behaviour is therefore needed in this novel ecological scenario where species populations are increasing and management needs to prepare and react.2.In this study, we investigated how densities of multiple herbivore species affect forest damage on commercially important Scots pine (Pinus sylvestris; pine) at the landscape scale across a latitudinal gradient. analysed multi-annual data from eight study landscapes to test the relationship between herbivore species densities and winter damage on pine (PWD) in young forest stands. As a density index, we used dung pellet group counts for herbivores of different body size and feeding types (moose, Alces alces; roe deer, Capreolus capreolus; red deer, Cervus elaphus; fallow deer, Dama dama). We applied the theory of associational effects to explore the relationship between PWD and the densities of pine and birch spp. at the forest stand scale, and also investigated the role of neighbourhood effects of the landscapes surrounding the forest stands on PWD,.3.Across the eight study areas, PWD increased with increasing moose density. Contrary to our expectations, higher densities of the smaller deer did not affect PWD significantly. Tree densities at the forest stand scale explained more variation in PWD than moose density. PWD decreased with increasing pine density and increased with increasing birch spp. density. This shows that densities of the target species important and suggests associational effects between pine and birch. Neighbouring land-use characteristics at the landscape scale did not affect PWD significantly.4.Synthesis and applications. We show consistent importance of forage abundance as well as associational effects in predicting the browsing patterns by large herbivores across a large landscape gradient. Our results suggest that for managers aiming to reduce PWD by deer, reducing deer densities alone might not be an ideal strategy. To control damage levels, we suggest that a combination of actions is needed. At the forest stand scale, plant-plant-interactions influencing associational effects need to be considered if PWD be reduced.

Hunting is a widespread but often overlooked land-use activity, providing major benefits to society. Hunting takes place in most landscapes, yet it remains unclear which types of landscapes foster or dampen hunting-related services, and how hunting relates to other land uses. A better understanding of these relationships is key for sustainable land-use planning that integrates wildlife management. This is particularly urgent for Europe, where wildlife populations are increasing. Focusing on Sweden, we explored the spatial associations among hunting, agriculture, and forestry to identify archetypical combinations of these land uses. Specifically, we combined indicators on the extent and intensity of agriculture and forestry, with data on hunting bags for 63 game species using self-organizing maps, a non-parametric clustering approach. We identified 15 typical bundles of co-occurring land uses at the municipality level across Sweden. The harvest of forest grouse, bears, and moose co-occurred with forestry in northern Sweden, whereas the harvest of small game, different deer species, and wild boar co-occurred with agriculture across southern Sweden, reflecting species' biology, environmental factors, and management. Our findings also highlight the strength of associations among hunting and other land uses. Importantly, we identified large areas in central Sweden where harvest of game was below average, possibly indicating that intensity of hunting is out of balance with that of agriculture or forestry, potentially fostering conflict between wildlife and land use. Collectively, our results suggest that (1) hunting should be considered a major land use that, in Sweden, is more widespread than agriculture and forestry; (2) land-use planning must therefore integrate wildlife management; and (3) such an integration should occur in a regionalized manner that considers social-ecological context. Our approach identifies a first spatial template within which such context-specific land-use planning, aiming at aligning wildlife and diverse land uses, can take place.