Synergistic effects of methyl jasmonate treatment and propagation method on Norway spruce resistance against a bark-feeding insect.

Utilizing plants with enhanced resistance traits is gaining interest in plant protection. Two strategies are especially promising for increasing resistance against a forest insect pest, the pine weevil (Hylobius abietis): exogenous application of the plant defense hormone methyl jasmonate (MeJA), and production of plants through the clonal propagation method somatic embryogenesis (SE). Here, we quantified and compared the separate and combined effects of SE and MeJA on Norway spruce resistance to pine weevil damage. Plants produced via SE (emblings) and nursery seedlings (containerized and bare-root), were treated (or not) with MeJA and exposed to pine weevils in the field (followed for 3 years) and in the lab (with a non-choice experiment). Firstly, we found that SE and MeJA independently decreased pine weevil damage to Norway spruce plants in the field by 32-33% and 53-59%, respectively, compared to untreated containerized and bare-root seedlings. Secondly, SE and MeJA together reduced damage to an even greater extent, with treated emblings receiving 86-87% less damage when compared to either untreated containerized or bare-root seedlings in the field, and by 48% in the lab. Moreover, MeJA-treated emblings experienced 98% lower mortality than untreated containerized seedlings, and this high level of survival was similar to that experienced by treated bare-root seedlings. These positive effects on survival remained for MeJA-treated emblings across the 3-year experimental period. We conclude that SE and MeJA have the potential to work synergistically to improve plants' ability to resist damage, and can thus confer a strong plant protection advantage. The mechanisms underlying these responses merit further examination.

Northward range expansion of rooting ungulates decreases detritivore and predatory mite abundances in boreal forests.

In the last few decades wild boar populations have expanded northwards, colonizing boreal forests. The soil disturbances caused by wild boar rooting may have an impact on soil organisms that play a key role in organic matter turnover. However, the impact of wild boar colonization on boreal forest ecosystems and soil organisms remains largely unknown. We investigated the effect of natural and simulated rooting on decomposer and predatory soil mites (total, adult and juvenile abundances; and adult-juvenile proportion). Our simulated rooting experiment aimed to disentangle the effects of (i) bioturbation due to soil mixing and (ii) removing organic material (wild boar food resources) on soil mites. Our results showed a decline in the abundance of adult soil mites in response to both natural and artificial rooting, while juvenile abundance and the relative proportion of adults and juveniles were not affected. The expansion of wild boar northwards and into new habitats has negative effects on soil decomposer abundances in boreal forests which may cascade through the soil food web ultimately affecting ecosystem processes. Our study also suggests that a combined use of natural and controlled experimental approaches is the way forward to reveal any subtle interaction between aboveground and belowground organisms and the ecosystem functions they drive.

Associating Land Cover Changes with Patterns of Incidences of Climate-Sensitive Infections: An Example on Tick-Borne Diseases in the Nordic Area.

Some of the climate-sensitive infections (CSIs) affecting humans are zoonotic vector-borne diseases, such as Lyme borreliosis (BOR) and tick-borne encephalitis (TBE), mostly linked to various species of ticks as vectors. Due to climate change, the geographical distribution of tick species, their hosts, and the prevalence of pathogens are likely to change. A recent increase in human incidences of these CSIs in the Nordic regions might indicate an expansion of the range of ticks and hosts, with vegetation changes acting as potential predictors linked to habitat suitability. In this paper, we study districts in Fennoscandia and Russia where incidences of BOR and TBE have steadily increased over the 1995-2015 period (defined as 'Well Increasing districts'). This selection is taken as a proxy for increasing the prevalence of tick-borne pathogens due to increased habitat suitability for ticks and hosts, thus simplifying the multiple factors that explain incidence variations. This approach allows vegetation types and strengths of correlation specific to the WI districts to be differentiated and compared with associations found over all districts. Land cover types and their changes found to be associated with increasing human disease incidence are described, indicating zones with potential future higher risk of these diseases. Combining vegetation cover and climate variables in regression models shows the interplay of biotic and abiotic factors linked to CSI incidences and identifies some differences between BOR and TBE. Regression model projections up until 2070 under different climate scenarios depict possible CSI progressions within the studied area and are consistent with the observed changes over the past 20 years.

Comparing Exogenous Methods to Induce Plant-Resistance Against a Bark-Feeding Insect.

Exogenous application of the plant hormone methyl jasmonate (MeJA) can trigger induced plant defenses against herbivores, and has been shown to provide protection against insect herbivory in conifer seedlings. Other methods, such as mechanical damage to seedlings, can also induce plant defenses, yet few have been compared to MeJA and most studies lack subsequent herbivory feeding tests. We conducted two lab experiments to: (1) compare the efficacy of MeJA to mechanical damage treatments that could also induce seedling resistance, (2) examine if subsequent insect damage differs depending on the time since induction treatments occurred, and (3) assess if these induction methods affect plant growth. We compared Scots pine (Pinus sylvestris) seedlings sprayed with MeJA (10 or 15 mM) to seedlings subjected to four different mechanical bark damage treatments (two different bark wound sizes, needle-piercing damage, root damage) and previous pine weevil (Hylobius abietis) damage as a reference treatment. The seedlings were exposed to pine weevils 12 or 32 days after treatments (early and late exposure, hereafter), and resistance was measured as the amount of damage received by plants. At early exposure, seedlings treated with needle-piercing damage received significantly more subsequent pine weevil feeding damage than those treated with MeJA. Seedlings treated with MeJA and needle-piercing damage received 84% less and 250% more pine weevil feeding, respectively, relative to control seedlings. The other treatments did not differ statistically from control or MeJA in terms of subsequent pine weevil damage. For the late exposure group, plants in all induction treatments tended to receive less pine weevil feeding (yet this was not statistically significant) compared to control seedlings. On the other hand, MeJA significantly slowed down seedling growth relative to control and all other induction treatments. Overall, the mechanical damage treatments appeared to have no or variable effects on seedling resistance. One of the treatments, needle-piercing damage, actually increased pine weevil feeding at early exposure. These results therefore suggest that mechanical damage shows little potential as a plant protection measure to reduce feeding by a bark-chewing insect.

Global change calls for novel plant protection: reviewing the potential of omnivorous plant-inhabiting arthropods as predators and plant defence inducers.

Global change poses new challenges for pest management. Omnivorous predatory arthropods play an important role in pest management, yet their potential has not been fully explored. Not only do they consume prey, but their plant-feeding induces plant defences that decrease herbivores' performance, and increases production of volatiles that attract natural enemies. Growing evidence from different plant-arthropod systems indicates the generality of plant defence induction following omnivore plant-feeding. Furthermore, these responses appear to affect other organisms (e.g. plant viruses), altering multi-trophic interactions. Here, we review the dual role of omnivores (as predators and plant inducers), identify knowledge gaps and provide future perspectives to increase our understanding of omnivores' multiple functions, and how this can be applied to advance plant protection strategies.

Interacting effects of insect and ungulate herbivory on Scots pine growth.

Most plants are subjected to damage from multiple species of herbivores, and the combined impact on plant growth can be non-additive. Since plant response to herbivores tends to be species specific, and change with repeated damage, the outcome likely depend on the sequence and number of attacks. There is a high likelihood of non-additive effects on plant growth by damage from mammals and insects, as mammalian herbivory can alter insect herbivore damage levels, yet few studies have explored this. We report the growth response of young Scots pine trees to sequential mammal and insect herbivory, varying the sequence and number of damage events, using an ungulate-pine-sawfly system. Combined sawfly and ungulate herbivory had both additive and non-additive effects on pine growth-the growth response depended on the combination of ungulate browsing and sawfly defoliation (significant interaction effect). Repeated sawfly herbivory reduced growth (compared to single defoliation) on un-browsed trees. However, on browsed trees, depending on when sawfly defoliation was combined with browsing, trees exposed to repeated sawfly herbivory had both higher, lower and the same growth as trees exposed to a single defoliation event. We conclude that the sequence of attacks by multiple herbivores determine plant growth response.

The tree species matters: Biodiversity and ecosystem service implications of replacing Scots pine production stands with Norway spruce.

The choice of tree species used in production forests matters for biodiversity and ecosystem services. In Sweden, damage to young production forests by large browsing herbivores is helping to drive a development where sites traditionally regenerated with Scots pine (Pinus sylvestris) are instead being regenerated with Norway spruce (Picea abies). We provide a condensed synthesis of the available evidence regarding the likely resultant implications for forest biodiversity and ecosystem services from this change in tree species. Apart from some benefits (e.g. reduced stand-level browsing damage), we identified a range of negative outcomes for biodiversity, production, esthetic and recreational values, as well as increased stand vulnerability to storm, frost, and drought damage, and potentially higher risks of pest and pathogen outbreak. Our results are directly relevant to forest owners and policy-makers seeking information regarding the uncertainties, risks, and trade-offs likely to result from changing the tree species in production forests.

Identifying climate-sensitive infectious diseases in animals and humans in Northern regions.

BACKGROUND: General knowledge on climate change effects and adaptation strategies has increased significantly in recent years. However, there is still a substantial information gap regarding the influence of climate change on infectious diseases and how these diseases should be identified. From a One Health perspective, zoonotic infections are of particular concern. The climate in Northern regions is changing faster than the global average. This study sought to identify climate-sensitive infectious diseases (CSIs) of relevance for humans and/or animals living in Northern regions. Inclusion criteria for CSIs were constructed using expert assessments. Based on these principles, 37 potential CSIs relevant for Northern regions were identified. A systematic literature search was performed in three databases using an explicit stepwise approach to determine whether the literature supports selection of these 37 potential CSIs. RESULTS: In total, 1275 nominated abstracts were read and categorised using predefined criteria. Results showed that arthropod vector-borne diseases in particular are recognised as having potential to expand their distribution towards Northern latitudes and that tick-borne encephalitis and borreliosis, midge-borne bluetongue and the parasitic infection fasciolosis can be classified as climate-sensitive. Many of the other potential CSIs considered are affected by extreme weather events, but could not be clearly classified as climate-sensitive. An additional literature search comparing awareness of climate influences on potential CSIs between 1997-2006 and 2007-2016 showed an increase in the number of papers mentioning effects of climate change. CONCLUSIONS: The four CSIs identified in this study could be targeted in a systematic surveillance programme in Northern regions. It is evident that climate change can affect the epidemiology and geographical range of many infectious diseases, but there were difficulties in identifying additional CSIs, most likely because other factors may be of equal or greater importance. However, climate-ecological dynamics are constantly under change, and therefore diseases may fall in or out of the climate-sensitive definition over time. There is increasing awareness in the literature of the effects of climate change on infectious diseases over time.

Tri-trophic interactions: bridging species, communities and ecosystems.

A vast body of research demonstrates that many ecological and evolutionary processes can only be understood from a tri-trophic viewpoint, that is, one that moves beyond the pairwise interactions of neighbouring trophic levels to consider the emergent features of interactions among multiple trophic levels. Despite its unifying potential, tri-trophic research has been fragmented, following two distinct paths. One has focused on the population biology and evolutionary ecology of simple food chains of interacting species. The other has focused on bottom-up and top-down controls over the distribution of biomass across trophic levels and other ecosystem-level variables. Here, we propose pathways to bridge these two long-standing perspectives. We argue that an expanded theory of tri-trophic interactions (TTIs) can unify our understanding of biological processes across scales and levels of organisation, ranging from species evolution and pairwise interactions to community structure and ecosystem function. To do so requires addressing how community structure and ecosystem function arise as emergent properties of component TTIs, and, in turn, how species traits and TTIs are shaped by the ecosystem processes and the abiotic environment in which they are embedded. We conclude that novel insights will come from applying tri-trophic theory systematically across all levels of biological organisation.

Trait-mediated indirect interactions: Moose browsing increases sawfly fecundity through plant-induced responses.

Induced responses in plants, initiated by herbivory, create potential for trait-mediated indirect interactions among herbivores. Responses to an initial herbivore may change a number of plant traits that subsequently alter ecological processes with additional herbivores. Although common, indirect interactions between taxonomically distant herbivores, such as mammals and insects, are less studied than between taxonomically related species (i.e., insect-insect). In terms of mammal-insect interactions, effects on insect numbers (e.g., density) are relatively well studied, whereas effects on performance (e.g., fecundity) are rarely explored. Moreover, few studies have explored mammal-insect interactions on coniferous plants.The aim of this study was to investigate the effect of mammalian induced responses on insect performance. We specifically investigated the effect of moose (Alces alces) browsing on Scots pine (Pinus sylvestris) and subsequent effects on sawfly (Neodiprion sertifer) performance.Sawfly larvae were reared on browsed, clipped, and unbrowsed control pine trees in a controlled field experiment. Afterward, cocoon weight was measured. Needle C:N ratio and di-terpene content were measured in response to browsing.Sawfly performance was enhanced on trees browsed by moose. Cocoon weight (proxy for fecundity) was 9 and 13% higher on browsed and clipped trees compared to unbrowsed trees. Cocoon weight was weakly related to needle C:N ratio, and browsed trees had lower a C:N ratio compared to unbrowsed trees. Needle di-terpene content, known to affect sawfly performance, was neither affected by the browsing treatments nor did it correlate with sawfly weight.We conclude that mammalian herbivory can affect insect herbivore performance, with potential consequences for ecological communities and with particular importance for insect population dynamics. The measured plant variables could not fully explain the effect on sawfly performance providing a starting point for the consideration of additional plant responses induced by mammalian browsing affecting insect performance.

Transient synchrony among populations of five foliage-feeding Lepidoptera.

Studies of transient population dynamics have largely focused on temporal changes in dynamical behaviour, such as the transition between periods of stability and instability. This study explores a related dynamic pattern, namely transient synchrony during a 49-year period among populations of five sympatric species of forest insects that share host tree resources. The long time series allows a more comprehensive exploration of transient synchrony patterns than most previous studies. Considerable variation existed in the dynamics of individual species, ranging from periodic to aperiodic. We used time-averaged methods to investigate long-term patterns of synchrony and time-localized methods to detect transient synchrony. We investigated transient patterns of synchrony between species and related these to the species' varying density dependence structures; even species with very different density dependence exhibited at least temporary periods of synchrony. Observed periods of interspecific synchrony may arise from interactions with host trees (e.g., induced host defences), interactions with shared natural enemies or shared impacts of environmental stochasticity. The transient nature of synchrony observed here raises questions both about the identity of synchronizing mechanisms and how these mechanisms interact with the endogenous dynamics of each species. We conclude that these patterns are the result of interspecific interactions that act only temporarily to synchronize populations, after which differences in the endogenous population dynamics among the species acts to desynchronize their dynamics.

General Public Acceptance of Forest Risk Management Strategies in Sweden: Comparing Three Approaches to Acceptability.

Global change calls for more active approaches to forest risk management. To avoid unforeseen backlashes, it is necessary to examine the general public's acceptance of the risk management strategies. By drawing on different theoretical approaches (threat and prevention, performance evaluations, and forest cognitions), the present study examines predictors of acceptability in the general public in three counties in Sweden (N = 1,026). As expected, appraisals of threat mediated the effect of threat awareness on belief in risk prevention, and when examining performance evaluations, trust in responsible actors influenced acceptability via procedural satisfaction. However, the threat and prevention approach and the performance evaluation approach only explained low levels of the variance in acceptability of the examined strategies. Nevertheless, stronger ecological forest values, and favoring broadleaved forests, were found to be important to the acceptability of proactively implementing a more diverse forest to meet the expected challenges associated with global climate change.

Costs and benefits of omnivore-mediated plant protection: effects of plant-feeding on Salix growth more detrimental than expected.

Predators can decrease herbivore damage to plants, and this is often assumed to be beneficial to plant growth/reproduction without actual quantification. Moreover, previous studies have been biased towards strict carnivores and neglected the role of omnivorous predators in prey-suppression. Here, we examined the costs (reduction in growth) and benefits (increase in growth) of enemy-mediated plant protection via the omnivorous (prey and plant-feeding) Orthotylus marginalis, relative to herbivory by a detrimental insect pest of Salix spp. plantations, the beetle Phratora vulgatissima. In a first experiment, we compared the cost of adult beetle versus omnivore nymph plant-feeding, and assessed the (non-) additive effects of the two types of damage. In a second experiment, we quantified the reduction in plant damage resulting from beetle-egg feeding by omnivorous nymphs and subsequent benefits to plants. We found that plant-feeding by omnivores negatively affected plant growth and this effect was similar to the cost imposed by beetle herbivory. Furthermore, simultaneous damage effects were additive and more detrimental than individual effects. While egg-predation by omnivore nymphs completely prevented beetle damage to plants, there was no difference in plant growth relative to only herbivore-damaged plants and growth was still reduced compared to control plants. Thus, despite herbivore suppression, there was no benefit to plant growth of omnivore-mediated plant protection and the negative effects of omnivore plant-feeding remained. These results are a first for an omnivorous enemy, and provide novel and timely insights on the underlying assumptions of tri-trophic associations and their use for biocontrol of insect pests.

Consumptive and nonconsumptive effect ratios depend on interaction between plant quality and hunting behavior of omnivorous predators.

Predators not only consume prey but exert nonconsumptive effects in form of scaring, consequently disturbing feeding or reproduction. However, how alternative food sources and hunting mode interactively affect consumptive and nonconsumptive effects with implications for prey fitness have not been addressed, impending functional understanding of such tritrophic interactions. With a herbivorous beetle, two omnivorous predatory bugs (plant sap as alternative food, contrasting hunting modes), and four willow genotypes (contrasting suitability for beetle/omnivore), we investigated direct and indirect effects of plant quality on the beetles key reproductive traits (oviposition rate, clutch size). Using combinations of either or both omnivores on different plant genotypes, we calculated the contribution of consumptive (eggs predated) and nonconsumptive (fewer eggs laid) effect on beetle fitness, including a prey density-independent measure (c:nc ratio). We found that larger clutches increase egg survival in presence of the omnivore not immediately consuming all eggs. However, rather than lowering mean, the beetles generally responded with a frequency shift toward smaller clutches. However, female beetles decreased mean and changed clutch size frequency with decreasing plant quality, therefore reducing intraspecific exploitative competition among larvae. More importantly, variation in host plant quality (to omnivore) led to nonconsumptive effects between one-third and twice as strong as the consumptive effects. Increased egg consumption on plants less suitable to the omnivore may therefore be accompanied by less searching and disturbing the beetle, representing a "cost" to the indirect plant defense in the form of a lower nonconsumptive effect. Many predators are omnivores and altering c:nc ratios (with egg retention as the most direct link to prey fitness) via plant quality and hunting behavior should be fundamental to advance ecological theory and applications. Furthermore, exploring modulation of fitness traits by bottom-up and top-down effects will help to explain how and why species aggregate.

Methods to identify the prey of invertebrate predators in terrestrial field studies.

Predation is an interaction during which an organism kills and feeds on another organism. Past and current interest in studying predation in terrestrial habitats has yielded a number of methods to assess invertebrate predation events in terrestrial ecosystems. We provide a decision tree to select appropriate methods for individual studies. For each method, we then present a short introduction, key examples for applications, advantages and disadvantages, and an outlook to future refinements. Video and, to a lesser extent, live observations are recommended in studies that address behavioral aspects of predator-prey interactions or focus on per capita predation rates. Cage studies are only appropriate for small predator species, but often suffer from a bias via cage effects. The use of prey baits or analyses of prey remains are cheaper than other methods and have the potential to provide per capita predation estimates. These advantages often come at the cost of low taxonomic specificity. Molecular methods provide reliable estimates at a fine level of taxonomic resolution and are free of observer bias for predator species of any size. However, the current PCR-based methods lack the ability to estimate predation rates for individual predators and are more expensive than other methods. Molecular and stable isotope analyses are best suited to address systems that include a range of predator and prey species. Our review of methods strongly suggests that while in many cases individual methods are sufficient to study specific questions, combinations of methods hold a high potential to provide more holistic insights into predation events. This review presents an overview of methods to researchers that are new to the field or to particular aspects of predation ecology and provides recommendations toward the subset of suitable methods to identify the prey of invertebrate predators in terrestrial field research.

Habitat heterogeneity affects predation of European pine sawfly cocoons.

Habitat heterogeneity is thought to affect top-down control of herbivorous insects and contribute to population stability by providing a more attractive microhabitat for natural enemies, potentially leading to reduced population fluctuations. Identifying the parameters that contribute to habitat heterogeneity promoting top-down control of herbivorous insects by natural enemies could facilitate appropriate management decisions, resulting in a decreased risk of pest insect outbreaks because of a higher level of predation. In our study, we measured the top-down pressure exerted by small mammals on the cocoons of a notorious pest insect in pine forests, the European pine sawfly (Neodiprion sertifer), which is known to be regulated by small mammal predation. The forest stands used differed in heterogeneity measured in terms of differences in tree diversity and density, understory vegetation height, presence/absence, and density of dead wood. We found higher predation in more dense spots within forest stands. Further, the effect of dead wood on sawfly cocoon predation depended on the pine proportion in forest stands. The addition of dead wood in a manipulation experiment had a slight positive effect on cocoon predation, while dead wood removal caused a clear decrease in predation rate, and the decrease was more pronounced when the proportion of pine increased. Our results show that habitat heterogeneity affects predation by generalist predators on herbivorous insects. This knowledge could be applied to reduce the risk of insect outbreaks by applying management methods that increase heterogeneity in perennial systems such as forests and orchards, thus decreasing the levels of insect damage.

Enhanced leaf nitrogen status stabilizes omnivore population density.

Plant traits can mediate the strength of interactions between omnivorous predators and their prey through density effects and changes in the omnivores' trophic behavior. In this study, we explored the established assumption that enhanced nutrient status in host plants strengthens the buffering effect of plant feeding for omnivorous predators, i.e., prevents rapid negative population growth during prey density decline and thereby increases and stabilizes omnivore population density. We analyzed 13 years of field data on population densities of a heteropteran omnivore on Salix cinerea stands, arranged along a measured leaf nitrogen gradient and found a 195 % increase in omnivore population density and a 63 % decrease in population variability with an increase in leaf nitrogen status from 26 to 40 mgN × g-1. We recreated the leaf nitrogen gradient in a greenhouse experiment and found, as expected, that increasing leaf nitrogen status enhanced omnivore performance but reduced per capita prey consumption. Feeding on high nitrogen status host plants can potentially decouple omnivore-prey population dynamics and allow omnivores to persist and function effectively at low prey densities to provide "background level" control of insect herbivores. This long-term effect is expected to outweigh the short-term effect on per capita prey consumption-resulting in a net increase in population predation rates with increasing leaf nitrogen status. Conservation biological control of insect pests that makes use of omnivore background control could, as a result, be manipulated via management of crop nitrogen status.

Female Salix viminalis are more severely infected by Melampsora spp. but neither sex experiences associational effects.

Associational effects of plant genotype or species on plant biotic interactions are common, not least for disease spread, but associational effects of plant sex on interactions have largely been ignored. Sex in dioecious plants can affect biotic interactions with herbivores and pollinators; however, its effects on plant-pathogen interactions are understudied and associational effects are unknown. In a replicated field experiment, we assessed Melampsora spp. leaf rust infection in monosexual and mixed sex plots of dioecious Salix viminalis L. to determine whether plant sex has either direct or associational effects on infection severity. We found no differences in Melampsora spp. infection severity among sexual monocultures and mixtures in our field experiment. However, female plants were overall more severely infected. In addition, we surveyed previous studies of infection in S. viminalis clones and reevaluated the studies after we assigned sex to the clones. We found that females were generally more severely infected, as in our field study. Similarly, in a survey of studies on sex-biased infection in dioecious plants, we found more female-biased infections in plant-pathogen pairs. We conclude that there was no evidence for associational plant sex effects of neighboring conspecifics for either females or males on infection severity. Instead, plant sex effects on infection act at an individual plant level. Our findings also suggest that female plants may in general be more severely affected by fungal pathogens than males.

Socio-ecological implications of modifying rotation lengths in forestry.

The rotation length is a key component of even-aged forest management systems. Using Fennoscandian forestry as a case, we review the socio-ecological implications of modifying rotation lengths relative to current practice by evaluating effects on a range of ecosystem services and on biodiversity conservation. The effects of shortening rotations on provisioning services are expected to be mostly negative to neutral (e.g. production of wood, bilberries, reindeer forage), while those of extending rotations would be more varied. Shortening rotations may help limit damage by some of today's major damaging agents (e.g. root rot, cambium-feeding insects), but may also increase other damage types (e.g. regeneration pests) and impede climate mitigation. Supporting (water, soil nutrients) and cultural (aesthetics, cultural heritage) ecosystem services would generally be affected negatively by shortened rotations and positively by extended rotations, as would most biodiversity indicators. Several effect modifiers, such as changes to thinning regimes, could alter these patterns.

Replacing monocultures with mixed-species stands: Ecosystem service implications of two production forest alternatives in Sweden.

Whereas there is evidence that mixed-species approaches to production forestry in general can provide positive outcomes relative to monocultures, it is less clear to what extent multiple benefits can be derived from specific mixed-species alternatives. To provide such insights requires evaluations of an encompassing suite of ecosystem services, biodiversity, and forest management considerations provided by specific mixtures and monocultures within a region. Here, we conduct such an assessment in Sweden by contrasting even-aged Norway spruce (Picea abies)-dominated stands, with mixed-species stands of spruce and birch (Betula pendula or B. pubescens), or spruce and Scots pine (Pinus sylvestris). By synthesizing the available evidence, we identify positive outcomes from mixtures including increased biodiversity, water quality, esthetic and recreational values, as well as reduced stand vulnerability to pest and pathogen damage. However, some uncertainties and risks were projected to increase, highlighting the importance of conducting comprehensive interdisciplinary evaluations when assessing the pros and cons of mixtures.