Enhancing multifunctionality in European boreal forests: The potential role of Triad landscape functional zoning.

Land-use policies aim at enhancing the sustainable use of natural resources. The Triad approach has been suggested to balance the social, ecological, and economic demands of forested landscapes. The core idea is to enhance multifunctionality at the landscape level by allocating landscape zones with specific management priorities, i.e., production (intensive management), multiple use (extensive management), and conservation (forest reserves). We tested the efficiency of the Triad approach and identified the respective proportion of above-mentioned zones needed to enhance multifunctionality in Finnish forest landscapes. Through a simulation and optimization framework, we explored a range of scenarios of the three zones and evaluated how changing their relative proportion (each ranging from 0 to 100%) impacted landscape multifunctionality, measured by various biodiversity and ecosystem service indicators. The results show that maximizing multifunctionality required around 20% forest area managed intensively, 50% extensively, and 30% allocated to forest reserves. In our case studies, such landscape zoning represented a good compromise between the studied multifunctionality components and maintained 61% of the maximum achievable net present value (i.e., total timber economic value). Allocating specific proportion of the landscape to a management zone had distinctive effects on the optimized economic or multifunctionality values. Net present value was only moderately impacted by shifting from intensive to extensive management, while multifunctionality benefited from less intensive and more diverse management regimes. This is the first study to apply Triad in a European boreal forest landscape, highlighting the usefulness of this approach. Our results show the potential of the Triad approach in promoting forest multifunctionality, as well as a strong trade-off between net present value and multifunctionality. We conclude that simply applying the Triad approach does not implicitly contribute to an overall increase in forest multifunctionality, as careful forest management planning still requires clear landscape objectives.

Increasing human environmental footprint does not lead to biotic homogenization of forest bird communities in northern USA.

Studies have shown negative impacts of increased human pressures on biodiversity at local (alpha-diversity) and regional (gamma-diversity) scales. However, the diversity between local sites (beta-diversity) has received less attention. This is an important shortcoming since beta-diversity acts as a linkage between the local and regional scales. Decreased beta-diversity means that local sites lose their distinctiveness, becoming more similar to each other. This process is known as biotic homogenization. However, the mechanisms causing biotic homogenization have not been fully studied nor its impacts on different facets of biodiversity. We examined if land-use change due to human actions causes biotic homogenization of taxonomic, functional, and phylogenetic diversity in bird communities of forested habitats in the state of Minnesota, USA. We address if forest loss and increased human domination in a region were associated with decreased beta-diversity. Our results showed that elevated human pressure was not related to increased biotic homogenization in this study region. Effects of landscape change were incongruent among taxonomic, functional, and phylogenetic diversity. At all spatial scales, taxonomic diversity was unrelated to forest loss or human domination. Interestingly, increased human domination appeared to increase the functional beta-diversity of bird communities. This association was driven by a decrease in local diversity. Forest habitat loss was associated with decreasing functional and phylogenetic diversity in local communities (alpha-diversity) and in regional species pool (gamma-diversity), but not in beta-diversity. We highlight the importance of considering multiple facets of biodiversity as their responses to human land-use is varied. Conservation significance of beta-diversity hinges on local and regional diversity responses to human land-use intensification, and organization of biodiversity should therefore be analyzed at multiple spatial scales.

Future supply of boreal forest ecosystem services is driven by management rather than by climate change.

Forests provide a wide variety of ecosystem services (ES) to society. The boreal biome is experiencing the highest rates of warming on the planet and increasing demand for forest products. To foresee how to maximize the adaptation of boreal forests to future warmer conditions and growing demands of forest products, we need a better understanding of the relative importance of forest management and climate change on the supply of ecosystem services. Here, using Finland as a boreal forest case study, we assessed the potential supply of a wide range of ES (timber, bilberry, cowberry, mushrooms, carbon storage, scenic beauty, species habitat availability and deadwood) given seven management regimes and four climate change scenarios. We used the forest simulator SIMO to project forest dynamics for 100 years into the future (2016-2116) and estimate the potential supply of each service using published models. Then, we tested the relative importance of management and climate change as drivers of the future supply of these services using generalized linear mixed models. Our results show that the effects of management on the future supply of these ES were, on average, 11 times higher than the effects of climate change across all services, but greatly differed among them (from 0.53 to 24 times higher for timber and cowberry, respectively). Notably, the importance of these drivers substantially differed among biogeographical zones within the boreal biome. The effects of climate change were 1.6 times higher in northern Finland than in southern Finland, whereas the effects of management were the opposite-they were three times higher in the south compared to the north. We conclude that new guidelines for adapting forests to global change should account for regional differences and the variation in the effects of climate change and management on different forest ES.

Breeding near heterospecifics as a defence against brood parasites: can redstarts lower probability of cuckoo parasitism using neighbours?

Breeding habitat choice based on the attraction to other species can provide valuable social information and protection benefits. In birds, species with overlapping resources can be a cue of good quality habitats; species with shared predators and/or brood parasites can increase joint vigilance or cooperative mobbing, while raptors may provide a protective umbrella against these threats. We tested whether the migratory common redstart (Phoenicurus phoenicurus) is attracted to breed near active nests of the great tit (Parus major), a keystone-information source for migrant passerine birds, or a top predator, the northern goshawk (Accipiter gentilis). This system is unique to test these questions because the redstart is a regular host for the common cuckoo (Cuculus canorus). Therefore, we also evaluated other possible benefits coming from the heterospecific attraction, especially in terms of reducing brood parasitism risk. We monitored redstart occupancy rates, onset of breeding, reproductive investment, and followed nest outcomes in terms of brood parasitism, nest predation risk and overall reproductive success. Redstarts avoided breeding near goshawks, but showed neither attraction nor avoidance to breed next to great tits. Both neighbours neither reduced brood parasitism risk nor affected overall nesting success in redstarts. Redstarts may not use heterospecific attraction for settlement decisions, as associations with other species can only exist when some benefits are gained. Thus, environmental cues may be more important than social information for redstarts when breeding habitat choice. Other front-line defence strategies may have a better impact reducing breeding negative interactions, such brood parasitism.

Titmice are a better indicator of bird density in Northern European than in Western European forests.

Population sizes of many birds are declining alarmingly and methods for estimating fluctuations in species' abundances at a large spatial scale are needed. The possibility to derive indicators from the tendency of specific species to co-occur with others has been overlooked. Here, we tested whether the abundance of resident titmice can act as a general ecological indicator of forest bird density in European forests. Titmice species are easily identifiable and have a wide distribution, which makes them potentially useful ecological indicators. Migratory birds often use information on the density of resident birds, such as titmice, as a cue for habitat selection. Thus, the density of residents may potentially affect community dynamics. We examined spatio-temporal variation in titmouse abundance and total bird abundance, each measured as biomass, by using long-term citizen science data on breeding forest birds in Finland and France. We analyzed the variation in observed forest bird density (excluding titmice) in relation to titmouse abundance. In Finland, forest bird density linearly increased with titmouse abundance. In France, forest bird density nonlinearly increased with titmouse abundance, the association weakening toward high titmouse abundance. We then analyzed whether the abundance (measured as biomass) of random species sets could predict forest bird density better than titmouse abundance. Random species sets outperformed titmice as an indicator of forest bird density only in 4.4% and 24.2% of the random draws, in Finland and France, respectively. Overall, the results suggest that titmice could act as an indicator of bird density in Northern European forest bird communities, encouraging the use of titmice observations by even less-experienced observers in citizen science monitoring of general forest bird density.

Forest management optimization across spatial scales to reconcile economic and conservation objectives.

Conflicts between biodiversity conservation and resource production can be mitigated by multi-objective management planning. Optimizing management for multiple objectives over larger land areas likely entails trading off the practicability of the process against the goodness of the solution. It is therefore worthwhile to resolve how large areas are required as management planning regions to reconcile conflicting objectives as effectively as possible. We aimed to reveal how the extent of forestry planning regions impacts the potential to mitigate a forestry-conservation conflict in Finland, represented as a trade-off between harvest income and deadwood availability. We used forecasted data from a forest simulator, a hierarchy of forestry planning regions, and an optimization model to explore the production possibility frontier between harvest income and deadwood. We compared the overall outcomes when management was optimized within the different-sized planning regions in terms of the two objectives, the spatial variation of deadwood, and the optimal combinations of management regimes. Increasing the size of the planning regions did produce higher simultaneous levels of the two objectives, but the differences were most often of the magnitude of only a few percentages. The differences among the scales were minor also in terms of the spatial variation in deadwood availability and in the optimal management combinations. The conflict between timber harvesting and deadwood availability is only marginally easier to mitigate at large spatial scales than at small forest ownership scales. However, regardless of the spatial scale of planning, the achievable solutions may not be good enough to safeguard deadwood-dependent biodiversity without active deadwood creation.

Interspecific information on predation risk affects nest site choice in a passerine bird.

BACKGROUND: Breeding site choice constitutes an important part of the species niche. Nest predation affects breeding site choice, and has been suggested to drive niche segregation and local coexistence of species. Interspecific social information use may, in turn, result in copying or rejection of heterospecific niche characteristics and thus affect realized niche overlap between species. We tested experimentally whether a migratory bird, the pied flycatcher Ficedula hypoleuca, collects information about nest predation risk from indirect cues of predators visiting nests of heterospecific birds. Furthermore, we investigated whether the migratory birds can associate such information with a specific nest site characteristic and generalize the information to their own nest site choice. RESULTS: Our results demonstrate that flycatchers can use the fate of heterospecific nesting attempts in their own nest site choice, but do so selectively. Young flycatcher females, when making the decision quickly, associated the fate of an artificial nest with nest-site characteristics and avoided the characteristic associated with higher nest predation risk. CONCLUSIONS: Copying nest site choices of successful heterospecifics, and avoiding choices which led to failed attempts, may amplify or counter effects of nest predation on niche overlap, with important consequences for between-species niche divergence-convergence dynamics, species coexistence and predator-prey interactions.

Environmental Characteristics and Anthropogenic Impact Jointly Modify Aquatic Macrophyte Species Diversity.

Species richness and spatial variation in community composition (i.e., beta diversity) are key measures of biodiversity. They are largely determined by natural factors, but also increasingly affected by anthropogenic factors. Thus, there is a need for a clear understanding of the human impact on species richness and beta diversity, the underlying mechanisms, and whether human-induced changes can override natural patterns. Here, we dissect the patterns of species richness, community composition and beta diversity in relation to different environmental factors as well as human impact in one framework: aquatic macrophytes in 66 boreal lakes in Eastern Finland. The lakes had been classified as having high, good or moderate status (according to ecological classification of surface waters in Finland) reflecting multifaceted human impact. We used generalized least square models to study the association between different environmental variables (Secchi depth, irregularity of the shoreline, total phosphorus, pH, alkalinity, conductivity) and species richness. We tested the null hypothesis that the observed community composition can be explained by random distribution of species. We used multivariate distance matrix regression to test the effect of each environmental variable on community composition, and distance-based test for homogeneity of multivariate dispersion to test whether lakes classified as high, good or moderate status have different beta diversity. We showed that environmental drivers of species richness and community composition were largely similar, although dependent on the particular life-form group studied. The most important ones were characteristics of water quality (pH, alkalinity, conductivity) and irregularity of the shoreline. Differences in community composition were related to environmental variables independently of species richness. Species richness was higher in lakes with higher levels of human impact. Lakes with different levels of human impact had different community composition. Between-lake beta diversity did not differ in high, good or moderate status groups. However, the variation in environmental variables shaping community composition was larger in lakes with moderate status compared to other lakes. Hence, beta diversity in lakes with moderate status was smaller than what could be expected on the basis of these environmental characteristics. This could be interpreted as homogenization.

Habitat quality is more important than matrix quality for bird communities in protected areas.

Protected areas are meant to preserve native local communities within their boundaries, but they are not independent from their surroundings. Impoverished habitat quality in the matrix might influence the species composition within the protected areas through biotic homogenization. The aim of this study was to determine the impacts of matrix quality on species richness and trait composition of bird communities from the Finnish reserve area network and whether the communities are being subject of biotic homogenization due to the lowered quality of the landscape matrix. We used joint species distribution modeling to study how characteristics of the Finnish forest reserves and the quality of their surrounding matrix alter species and trait compositions of forest birds. The proportion of old forest within the reserves was the main factor in explaining the bird community composition, and the bird communities within the reserves did not strongly depend on the quality of the matrix. Yet, in line with the homogenization theory, the beta-diversity within reserves embedded in low-quality matrix was lower than that in high-quality matrix, and the average abundance of regionally abundant species was higher. Influence of habitat quality on bird community composition was largely explained by the species' functional traits. Most importantly, the community specialization index was low, and average body size was high in areas with low proportion of old forest. We conclude that for conserving local bird communities in northern Finnish protected forests, it is currently more important to improve or maintain habitat quality within the reserves than in the surrounding matrix. Nevertheless, we found signals of bird community homogenization, and thus, activities that decrease the quality of the matrix are a threat for bird communities.

Degradation in landscape matrix has diverse impacts on diversity in protected areas.

INTRODUCTION: A main goal of protected areas is to maintain species diversity and the integrity of biological assemblages. Intensifying land use in the matrix surrounding protected areas creates a challenge for biodiversity conservation. Earlier studies have mainly focused on taxonomic diversity within protected areas. However, functional and especially phylogenetic diversities are less studied phenomena, especially with respect to the impacts of the matrix that surrounds protected areas. Phylogenetic diversity refers to the range of evolutionary lineages, the maintenance of which ensures that future evolutionary potential is safeguarded. Functional diversity refers to the range of ecological roles that members of a community perform. For ecosystem functioning and long-term resilience, they are at least as important as taxonomic diversity. AIM: We studied how the characteristics of protected areas and land use intensity in the surrounding matrix affect the diversity of bird communities in protected boreal forests. We used line-transect count and land-cover data from 91 forest reserves in Northern Finland, and land-cover data from buffer zones surrounding these reserves. We studied if habitat diversity and productivity inside protected areas, and intensity of forest management in the matrix have consistent effects on taxonomic, functional and phylogenetic diversities, and community specialization. RESULTS: We found that habitat diversity and productivity inside protected areas have strong effects on all diversity metrics, but matrix effects were inconsistent. The proportion of old forest in the matrix, reflecting low intensity forest management, had positive effects on community specialization. Interestingly, functional diversity increased with increasing logging intensity in the matrix. CONCLUSIONS: Our results indicate that boreal forest reserves are not able to maintain their species composition and abundances if embedded in a severely degraded matrix. Our study also highlights the importance of focusing on different aspects of biodiversity.

Impacts of forestry on boreal forests: An ecosystem services perspective.

Forests are widely recognized as major providers of ecosystem services, including timber, other forest products, recreation, regulation of water, soil and air quality, and climate change mitigation. Extensive tracts of boreal forests are actively managed for timber production, but actions aimed at increasing timber yields also affect other forest functions and services. Here, we present an overview of the environmental impacts of forest management from the perspective of ecosystem services. We show how prevailing forestry practices may have substantial but diverse effects on the various ecosystem services provided by boreal forests. Several aspects of these processes remain poorly known and warrant a greater role in future studies, including the role of community structure. Conflicts among different interests related to boreal forests are most likely to occur, but the concept of ecosystem services may provide a useful framework for identifying and resolving these conflicts.

Projecting biodiversity and wood production in future forest landscapes: 15 key modeling considerations.

A variety of modeling approaches can be used to project the future development of forest systems, and help to assess the implications of different management alternatives for biodiversity and ecosystem services. This diversity of approaches does however present both an opportunity and an obstacle for those trying to decide which modeling technique to apply, and interpreting the management implications of model output. Furthermore, the breadth of issues relevant to addressing key questions related to forest ecology, conservation biology, silviculture, economics, requires insights stemming from a number of distinct scientific disciplines. As forest planners, conservation ecologists, ecological economists and silviculturalists, experienced with modeling trade-offs and synergies between biodiversity and wood biomass production, we identified fifteen key considerations relevant to assessing the pros and cons of alternative modeling approaches. Specifically we identified key considerations linked to study question formulation, modeling forest dynamics, forest processes, study landscapes, spatial and temporal aspects, and the key response metrics - biodiversity and wood biomass production, as well as dealing with trade-offs and uncertainties. We also provide illustrative examples from the modeling literature stemming from the key considerations assessed. We use our findings to reiterate the need for explicitly addressing and conveying the limitations and uncertainties of any modeling approach taken, and the need for interdisciplinary research efforts when addressing the conservation of biodiversity and sustainable use of environmental resources.

Optimal conservation resource allocation under variable economic and ecological time discounting rates in boreal forest.

Resource allocation to multiple alternative conservation actions is a complex task. A common trade-off occurs between protection of smaller, expensive, high-quality areas versus larger, cheaper, partially degraded areas. We investigate optimal allocation into three actions in boreal forest: current standard forest management rules, setting aside of mature stands, or setting aside of clear-cuts. We first estimated how habitat availability for focal indicator species and economic returns from timber harvesting develop through time as a function of forest type and action chosen. We then developed an optimal resource allocation by accounting for budget size and habitat availability of indicator species in different forest types. We also accounted for the perspective adopted towards sustainability, modeled via temporal preference and economic and ecological time discounting. Controversially, we found that in boreal forest set-aside followed by protection of clear-cuts can become a winning cost-effective strategy when accounting for habitat requirements of multiple species, long planning horizon, and limited budget. It is particularly effective when adopting a long-term sustainability perspective, and accounting for present revenues from timber harvesting. The present analysis assesses the cost-effective conditions to allocate resources into an inexpensive conservation strategy that nevertheless has potential to produce high ecological values in the future.

The role of novel forest ecosystems in the conservation of wood-inhabiting fungi in boreal broadleaved forests.

The increasing human impact on the earth's biosphere is inflicting changes at all spatial scales. As well as deterioration and fragmentation of natural biological systems, these changes also led to other, unprecedented effects and emergence of novel habitats. In boreal zone, intensive forest management has negatively impacted a multitude of deadwood-associated species. This is especially alarming given the important role wood-inhabiting fungi have in the natural decay processes. In the boreal zone, natural broad-leaved-dominated, herb-rich forests are threatened habitats which have high wood-inhabiting fungal species richness. Fungal diversity in other broadleaved forest habitat types is poorly known. Traditional wood pastures and man-made afforested fields are novel habitats that could potentially be important for wood-inhabiting fungi. This study compares species richness and fungal community composition across the aforementioned habitat types, based on data collected for wood-inhabiting fungi occupying all deadwood diameter fractions. Corticioid and polyporoid fungi were surveyed from 67 130 deadwood particles in four natural herb-rich forests, four birch-dominated wood pastures, and four birch-dominated afforested field sites in central Finland. As predicted, natural herb-rich forests were the most species-rich habitat. However, afforested fields also had considerably higher overall species richness than wood pastures. Many rare or rarely collected species were detected in each forest type. Finally, fungal community composition showed some divergence not only among the different habitat types, but also among deadwood diameter fractions. Synthesis and applications: In order to maintain biodiversity at both local and regional scales, conserving threatened natural habitat types and managing traditional landscapes is essential. Man-made secondary woody habitats could provide the necessary resources and serve as surrogate habitats for many broadleaved deadwood-associated species, and thus complement the existing conservation network of natural forests.

Applying a framework for landscape planning under climate change for the conservation of biodiversity in the Finnish boreal forest.

Conservation strategies are often established without consideration of the impact of climate change. However, this impact is expected to threaten species and ecosystem persistence and to have dramatic effects towards the end of the 21st century. Landscape suitability for species under climate change is determined by several interacting factors including dispersal and human land use. Designing effective conservation strategies at regional scales to improve landscape suitability requires measuring the vulnerabilities of specific regions to climate change and determining their conservation capacities. Although methods for defining vulnerability categories are available, methods for doing this in a systematic, cost-effective way have not been identified. Here, we use an ecosystem model to define the potential resilience of the Finnish forest landscape by relating its current conservation capacity to its vulnerability to climate change. In applying this framework, we take into account the responses to climate change of a broad range of red-listed species with different niche requirements. This framework allowed us to identify four categories in which representation in the landscape varies among three IPCC emission scenarios (B1, low; A1B, intermediate; A2, high emissions): (i) susceptible (B1 = 24.7%, A1B = 26.4%, A2 = 26.2%), the most intact forest landscapes vulnerable to climate change, requiring management for heterogeneity and resilience; (ii) resilient (B1 = 2.2%, A1B = 0.5%, A2 = 0.6%), intact areas with low vulnerability that represent potential climate refugia and require conservation capacity maintenance; (iii) resistant (B1 = 6.7%, A1B = 0.8%, A2 = 1.1%), landscapes with low current conservation capacity and low vulnerability that are suitable for restoration projects; (iv) sensitive (B1 = 66.4%, A1B = 72.3%, A2 = 72.0%), low conservation capacity landscapes that are vulnerable and for which alternative conservation measures are required depending on the intensity of climate change. Our results indicate that the Finnish landscape is likely to be dominated by a very high proportion of sensitive and susceptible forest patches, thereby increasing uncertainty for landscape managers in the choice of conservation strategies.

Spatially dynamic forest management to sustain biodiversity and economic returns.

Production of marketed commodities and protection of biodiversity in natural systems often conflict and thus the continuously expanding human needs for more goods and benefits from global ecosystems urgently calls for strategies to resolve this conflict. In this paper, we addressed what is the potential of a forest landscape to simultaneously produce habitats for species and economic returns, and how the conflict between habitat availability and timber production varies among taxa. Secondly, we aimed at revealing an optimal combination of management regimes that maximizes habitat availability for given levels of economic returns. We used multi-objective optimization tools to analyze data from a boreal forest landscape consisting of about 30,000 forest stands simulated 50 years into future. We included seven alternative management regimes, spanning from the recommended intensive forest management regime to complete set-aside of stands (protection), and ten different taxa representing a wide variety of habitat associations and social values. Our results demonstrate it is possible to achieve large improvements in habitat availability with little loss in economic returns. In general, providing dead-wood associated species with more habitats tended to be more expensive than providing requirements for other species. No management regime alone maximized habitat availability for the species, and systematic use of any single management regime resulted in considerable reductions in economic returns. Compared with an optimal combination of management regimes, a consistent application of the recommended management regime would result in 5% reduction in economic returns and up to 270% reduction in habitat availability. Thus, for all taxa a combination of management regimes was required to achieve the optimum. Refraining from silvicultural thinnings on a proportion of stands should be considered as a cost-effective management in commercial forests to reconcile the conflict between economic returns and habitat required by species associated with dead-wood. In general, a viable strategy to maintain biodiversity in production landscapes would be to diversify management regimes. Our results emphasize the importance of careful landscape level forest management planning because optimal combinations of management regimes were taxon-specific. For cost-efficiency, the results call for balanced and correctly targeted strategies among habitat types.

Manipulating individual decisions and environmental conditions reveal individual quality in decision-making and non-lethal costs of predation risk.

Habitat selection is a crucial decision for any organism. Selecting a high quality site will positively impact survival and reproductive output. Predation risk is an important component of habitat quality that is known to impact reproductive success and individual condition. However, separating the breeding consequences of decision-making of wild animals from individual quality is difficult. Individuals face reproductive decisions that often vary with quality such that low quality individuals invest less. This reduced reproductive performance could appear a cost of increased risk but may simply reflect lower quality. Thus, teasing apart the effects of individual quality and the effect of predation risk is vital to understand the physiological and reproductive costs of predation risk alone on breeding animals. In this study we alter the actual territory location decisions of pied flycatchers by moving active nests relative to breeding sparrowhawks, the main predators of adult flycatchers. We experimentally measure the non-lethal effects of predation on adults and offspring while controlling for effects of parental quality, individual territory choice and initiation of breeding. We found that chicks from high predation risk nests (<50 m of hawk) were significantly smaller than chicks from low risk nests (>200 m from hawk). However, in contrast to correlative results, females in manipulated high risk nests did not suffer decreased body condition or increased stress response (HSP60 and HSP70). Our results suggest that territory location decisions relative to breeding avian predators cause spatial gradients in individual quality. Small adjustments in territory location decisions have crucial consequences and our results confirm non-lethal costs of predation risk that were expressed in terms of smaller offspring produced. However, females did not show costs in physiological condition which suggests that part of the costs incurred by adults exposed to predation risk are quality determined.

New behavioural trait adopted or rejected by observing heterospecific tutor fitness.

Animals can acquire behaviours from others, including heterospecifics, but should be discriminating in when and whom to copy. Successful individuals should be preferred as tutors, while adopting traits of poorly performing individuals should be actively avoided. Thus far it is unknown if such adaptive strategies are involved when individuals copy other species. Furthermore, rejection of traits based on tutor characteristics (negative bias) has not been shown in any non-human animal. Here we test whether a choice between two new, neutral behavioural alternatives-breeding-sites with alternative geometric symbols-is affected by observing the choice and fitness of a heterospecific tutor. A field experiment replicated in four different areas shows that the proportion of pied flycatcher females matching the choice of the tit tutor consistently increased with increasing number of offspring in the tit nest, to the extent of nearly complete prevalence in one of the areas when tit fitness was highest. Notably, all four replicates demonstrate rejection of the behaviour of lowest-fitness tutors. The results demonstrate both acquisition and avoidance of heterospecific behavioural traits, based on the perceived (lack of) tutor fitness. This has potential implications for understanding the origin, diversity and local adaptations of behavioural traits, and niche overlap/partitioning and species co-occurrence.