Ecological and evolutionary consequences of selective interspecific information use.

Recent work has shown that animals frequently use social information from individuals of their own species as well as from other species; however, the ecological and evolutionary consequences of this social information use remain poorly understood. Additionally, information users may be selective in their social information use, deciding from whom and how to use information, but this has been overlooked in an interspecific context. In particular, the intentional decision to reject a behaviour observed via social information has received less attention, although recent work has indicated its presence in various taxa. Based on existing literature, we explore in which circumstances selective interspecific information use may lead to different ecological and coevolutionary outcomes between two species, such as explaining observed co-occurrences of putative competitors. The initial ecological differences and the balance between the costs of competition and the benefits of social information use potentially determine whether selection may lead to trait divergence, convergence or coevolutionary arms race between two species. We propose that selective social information use, including adoption and rejection of behaviours, may have far-reaching fitness consequences, potentially leading to community-level eco-evolutionary outcomes. We argue that these consequences of selective interspecific information use may be much more widespread than has thus far been considered.

Size of an interspecific competitor may be a source of information in reproductive decisions.

Animals use inter-specific cues as a source of information in decisions-making, but the full costs and benefits of inter-specific information use are unknown. We tested whether pied flycatchers use the body size and clutch size of great tits as cues in their reproductive decisions and what are the possible fitness consequences as a function of great tit size. The size of great tit females associated positively with flycatcher's probability to settle near a tit nest over a territory further away. Flycatcher egg mass was positively correlated with great tit female size regardless of flycatcher territory choice. However, in flycatchers that had chosen to nest near great tits, the size of nestlings decreased in relation to increasing great tit female size. Our results demonstrate the use of size of inter-specifics as a cue in reproductive decisions and the trade-off between the value of information and costs of competition information users face when using inter-specific information in decision-making.

Urban forest soils harbour distinct and more diverse communities of bacteria and fungi compared to less disturbed forest soils.

Anthropogenic changes to land use drive concomitant changes in biodiversity, including that of the soil microbiota. However, it is not clear how increasing intensity of human disturbance is reflected in the soil microbial communities. To address this issue, we used amplicon sequencing to quantify the microbiota (bacteria and fungi) in the soil of forests (n = 312) experiencing four different land uses, national parks (set aside for nature conservation), managed (for forestry purposes), suburban (on the border of an urban area) and urban (fully within a town or city), which broadly represent a gradient of anthropogenic disturbance. Alpha diversity of bacteria and fungi increased with increasing levels of anthropogenic disturbance, and was thus highest in urban forest soils and lowest in the national parks. The forest soil microbial communities were structured according to the level of anthropogenic disturbance, with a clear urban signature evident in both bacteria and fungi. Despite notable differences in community composition, there was little change in the predicted functional traits of urban bacteria. By contrast, urban soils exhibited a marked loss of ectomycorrhizal fungi. Soil pH was positively correlated with the level of disturbance, and thus was the strongest predictor of variation in alpha and beta diversity of forest soil communities, indicating a role of soil alkalinity in structuring urban soil microbial communities. Hence, our study shows how the properties of urban forest soils promote an increase in microbial diversity and a change in forest soil microbiota composition.

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.

Conceptual preferences can be transmitted via selective social information use between competing wild bird species.

Concept learning is considered a high-level adaptive ability. Thus far, it has been studied in laboratory via asocial trial and error learning. Yet, social information use is common among animals but it remains unknown whether concept learning by observing others occurs. We tested whether pied flycatchers (Ficedula hypoleuca) form conceptual relationships from the apparent choices of nest-site characteristics (geometric symbol attached to the nest-box) of great tits (Parus major). Each wild flycatcher female (n = 124) observed one tit pair that exhibited an apparent preference for either a large or a small symbol and was then allowed to choose between two nest-boxes with a large and a small symbol, but the symbol shape was different to that on the tit nest. Older flycatcher females were more likely to copy the symbol size preference of tits than yearling flycatcher females when there was a high number of visible eggs or a few partially visible eggs in the tit nest. However, this depended on the phenotype, copying switched to rejection as a function of increasing body size. Possibly the quality of and overlap in resource use with the tits affected flycatchers' decisions. Hence, our results suggest that conceptual preferences can be horizontally transmitted across coexisting animals, which may increase the performance of individuals that use concept learning abilities in their decision-making.

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.

Calf/female ratio and population dynamics of wild forest reindeer in relation to wolf and moose abundances in a managed European ecosystem.

BACKGROUND: The alternative prey hypothesis describes the mechanism for apparent competition whereby the mortality of the secondary prey species increases (and population size decreases decreases) by the increased predation by the shared predator if the population size of the primary prey decreases. Apparent competition is a process where the abundance of two co-existing prey species are negatively associated because they share a mutual predator, which negatively affects the abundance of both prey Here, we examined whether alternative prey and/or apparent competition hypothesis can explain the population dynamics and reproductive output of the secondary prey, wild forest reindeer (Rangifer tarandus fennicus) in Finland, in a predator-prey community in which moose (Alces alces) is the primary prey and the wolf (Canis lupus) is the generalist predator. METHODS: We examined a 22-year time series (1996-2017) to determine how the population size and the calf/female ratio of wild forest reindeer in Eastern Finland were related to the abundances of wolf and moose. Only moose population size was regulated by hunting. Summer predation of wolves on reindeer focuses on calves. We used least squares regression (GLS) models (for handling autocorrelated error structures and resulting pseudo-R2s) and generalized linear mixed (GLMs) models (for avoidance of negative predictions) to determine the relationships between abundances. We performed linear and general linear models for the calf/female ratio of reindeer. RESULTS AND SYNTHESIS: The trends in reindeer population size and moose abundance were almost identical: an increase during the first years and then a decrease until the last years of our study period. Wolf population size in turn did not show long-term trends. Change in reindeer population size between consecutive winters was related positively to the calf/female ratio. The calf/female ratio was negatively related to wolf population size, but the reindeer population size was related to the wolf population only when moose abundance was entered as another independent variable. The wolf population was not related to moose abundance even though it is likely to consist the majority of the prey biomass. Because reindeer and moose populations were positively associated, our results seemed to support the alternative prey hypothesis more than the apparent competition hypothesis. However, these two hypotheses are not mutually exclusive and the primary mechanism is difficult to distinguish as the system is heavily managed by moose hunting. The recovery of wild forest reindeer in eastern Finland probably requires ecosystem management involving both habitat restoration and control of species abundances.

Rodent host population dynamics drive zoonotic Lyme Borreliosis and Orthohantavirus infections in humans in Northern Europe.

Zoonotic diseases, caused by pathogens transmitted between other vertebrate animals and humans, pose a major risk to human health. Rodents are important reservoir hosts for many zoonotic pathogens, and rodent population dynamics affect the infection dynamics of rodent-borne diseases, such as diseases caused by hantaviruses. However, the role of rodent population dynamics in determining the infection dynamics of rodent-associated tick-borne diseases, such as Lyme borreliosis (LB), caused by Borrelia burgdorferi sensu lato bacteria, have gained limited attention in Northern Europe, despite the multiannual abundance fluctuations, the so-called vole cycles, that characterise rodent population dynamics in the region. Here, we quantify the associations between rodent abundance and LB human cases and Puumala Orthohantavirus (PUUV) infections by using two time series (25-year and 9-year) in Finland. Both bank vole (Myodes glareolus) abundance as well as LB and PUUV infection incidence in humans showed approximately 3-year cycles. Without vector transmitted PUUV infections followed the bank vole host abundance fluctuations with two-month time lag, whereas tick-transmitted LB was associated with bank vole abundance ca. 12 and 24 months earlier. However, the strength of association between LB incidence and bank vole abundance ca. 12 months before varied over the study years. This study highlights that the human risk to acquire rodent-borne pathogens, as well as rodent-associated tick-borne pathogens is associated with the vole cycles in Northern Fennoscandia, yet with complex time lags.

Quantitative genetics of the use of conspecific and heterospecific social cues for breeding site choice.

Social information use for decision-making is common and affects ecological and evolutionary processes, including social aggregation, species coexistence, and cultural evolution. Despite increasing ecological knowledge on social information use, very little is known about its genetic basis and therefore its evolutionary potential. Genetic variation in a trait affecting an individual's social and nonsocial environment may have important implications for population dynamics, interspecific interactions, and, for expression of other, environmentally plastic traits. We estimated repeatability, additive genetic variance, and heritability of the use of conspecific and heterospecific social cues (abundance and breeding success) for breeding site choice in a population of wild collared flycatchers Ficedula albicollis. Repeatability was found for two social cues: previous year conspecific breeding success and previous year heterospecific abundance. Yet, additive genetic variances for these two social cues, and thus heritabilities, were low. This suggests that most of the phenotypic variation in the use of social cues and resulting conspecific and heterospecific social environment experienced by individuals in this population stems from phenotypic plasticity. Given the important role of social information use on ecological and evolutionary processes, more studies on genetic versus environmental determinism of social information use are needed.

Behavioural traits modulate the use of heterospecific social information for nest site selection: experimental evidence from a wild bird population.

The use of social information for making decisions is common but can be constrained by behavioural traits via, for example, the ability to gather information. Such constrained information use has been described in foraging habitat selection; yet it remains unexplored in the breeding habitat selection context, despite potentially strong fitness consequences. We experimentally tested whether three behavioural traits (aggressiveness, boldness and neophobia) affected the use of heterospecific social information for nest site selection in wild collared flycatchers Ficedula albicollis. Flycatchers have previously been found to copy or reject an artificial apparent preference of tits (their main competitors) for a nest site feature: they preferred nest-boxes with the same or a different feature, depending on tit early reproductive investment. Here, we confirmed this result and showed that shy individuals and less aggressive old males (i.e. 2 years old or older) copied tit apparent preference, while more aggressive old males rejected the tit preference. Aggressiveness and boldness may allow males to access more information sources or affect males' interactions with dominant tits when selecting a nest site. Our study highlights the links between variation in behaviours and social information use for breeding habitat selection and calls for further work to explore underlying mechanisms.

Information collected during the post-breeding season guides future breeding decisions in a migratory bird.

Breeding habitat choice and investment decisions are key contributors to fitness in animals. Density of individuals is a well-known cue of habitat quality used for future breeding decisions, but accuracy of density cues decreases as individuals disperse from breeding sites. Used nests remain an available information source also after breeding season, but whether such information is used for breeding decisions is less well known. We experimentally investigated whether migratory, cavity-nesting pied flycatchers (Ficedula hypoleuca) prospect potential breeding sites after breeding season and use old nests as a cue for future breeding decisions. In late summer 2013, forest sites were assigned to four treatments: (1) sites including nest boxes with old nests of heterospecifics (tits), (2) sites including suitable but empty nest boxes, (3) sites with unsuitable nest boxes, or (4) sites without any nest boxes. In the following year, we investigated pied flycatcher habitat choice and reproductive investment according to these "past" cues while also controlling for additional information sources present during settlement. Flycatchers preferred sites where tits had been perceived to breed in the previous year, but only if great tits were also currently breeding in the site and had a relatively high number of eggs. Old flycatchers avoided sites previously treated with suitable but empty cavities, whereas young flycatchers preferred sites where tits had apparently bred in the previous year. Also egg mass, but not clutch size or clutch mass, was affected by the combination of past treatment information and current tit abundance.

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.

Interspecific variation in the relationship between clutch size, laying date and intensity of urbanization in four species of hole-nesting birds.

The increase in size of human populations in urban and agricultural areas has resulted in considerable habitat conversion globally. Such anthropogenic areas have specific environmental characteristics, which influence the physiology, life history, and population dynamics of plants and animals. For example, the date of bud burst is advanced in urban compared to nearby natural areas. In some birds, breeding success is determined by synchrony between timing of breeding and peak food abundance. Pertinently, caterpillars are an important food source for the nestlings of many bird species, and their abundance is influenced by environmental factors such as temperature and date of bud burst. Higher temperatures and advanced date of bud burst in urban areas could advance peak caterpillar abundance and thus affect breeding phenology of birds. In order to test whether laying date advance and clutch sizes decrease with the intensity of urbanization, we analyzed the timing of breeding and clutch size in relation to intensity of urbanization as a measure of human impact in 199 nest box plots across Europe, North Africa, and the Middle East (i.e., the Western Palearctic) for four species of hole-nesters: blue tits (Cyanistes caeruleus), great tits (Parus major), collared flycatchers (Ficedula albicollis), and pied flycatchers (Ficedula hypoleuca). Meanwhile, we estimated the intensity of urbanization as the density of buildings surrounding study plots measured on orthophotographs. For the four study species, the intensity of urbanization was not correlated with laying date. Clutch size in blue and great tits does not seem affected by the intensity of urbanization, while in collared and pied flycatchers it decreased with increasing intensity of urbanization. This is the first large-scale study showing a species-specific major correlation between intensity of urbanization and the ecology of breeding. The underlying mechanisms for the relationships between life history and urbanization remain to be determined. We propose that effects of food abundance or quality, temperature, noise, pollution, or disturbance by humans may on their own or in combination affect laying date and/or clutch size.

Variation in clutch size in relation to nest size in birds.

Nests are structures built to support and protect eggs and/or offspring from predators, parasites, and adverse weather conditions. Nests are mainly constructed prior to egg laying, meaning that parent birds must make decisions about nest site choice and nest building behavior before the start of egg-laying. Parent birds should be selected to choose nest sites and to build optimally sized nests, yet our current understanding of clutch size-nest size relationships is limited to small-scale studies performed over short time periods. Here, we quantified the relationship between clutch size and nest size, using an exhaustive database of 116 slope estimates based on 17,472 nests of 21 species of hole and non-hole-nesting birds. There was a significant, positive relationship between clutch size and the base area of the nest box or the nest, and this relationship did not differ significantly between open nesting and hole-nesting species. The slope of the relationship showed significant intraspecific and interspecific heterogeneity among four species of secondary hole-nesting species, but also among all 116 slope estimates. The estimated relationship between clutch size and nest box base area in study sites with more than a single size of nest box was not significantly different from the relationship using studies with only a single size of nest box. The slope of the relationship between clutch size and nest base area in different species of birds was significantly negatively related to minimum base area, and less so to maximum base area in a given study. These findings are consistent with the hypothesis that bird species have a general reaction norm reflecting the relationship between nest size and clutch size. Further, they suggest that scientists may influence the clutch size decisions of hole-nesting birds through the provisioning of nest boxes of varying sizes.

Avoiding perceived past resource use of potential competitors affects niche dynamics in a bird community.

BACKGROUND: Social information use is usually considered to lead to ecological convergence among involved con- or heterospecific individuals. However, recent results demonstrate that observers can also actively avoid behaving as those individuals being observed, leading to ecological divergence. This phenomenon has been little explored so far, yet it can have significant impact on resource use, realized niches and species co-existence. In particular, the time-scale and the ecological context over which such shifts can occur are unknown. We examined with a long-term (four years) field experiment whether experimentally manipulated, species-specific, nest-site feature preferences (symbols on nest boxes) are transmitted across breeding seasons and affect future nest-site preferences in a guild of three cavity-nesting birds. RESULTS: Of the examined species, resident great tits (Parus major) preferred the symbol that had been associated with unoccupied nest boxes in the previous year, i.e., their preference shifted towards niche space previously unused by putative competitors and conspecifics. CONCLUSIONS: Our results show that animals can remember the earlier resource use of conspecifics and other guild members and adjust own decisions accordingly one year after. Our experiment cannot reveal the ultimate mechanism(s) behind the observed behaviour but avoiding costs of intra- or interspecific competition or ectoparasite load in old nests are plausible reasons. Our findings imply that interspecific social information use can affect resource sharing and realized niches in ecological time-scale through active avoidance of observed decisions and behavior of potentially competing species.

Active hiding of social information from information-parasites.

BACKGROUND: Coevolution between pairs of different kind of entities, such as providers and users of information, involves reciprocal selection pressures between them as a consequence of their ecological interaction. Pied flycatchers (Ficedula hypoleuca) have been shown to derive fitness benefits (larger clutches) when nesting in proximity to great tits (Parus major), presumably because they this way discover and obtain information about nesting sites. Tits suffer from the resulting association (smaller clutches). An arms race between the tits (information host) and the flycatchers (information parasite) could thus result. Great tits often cover eggs with nesting material before, but not during incubation. We hypothesized that one function of egg-covering could be a counter-adaptation to reduce information parasitism by pied flycatchers. We predicted that tits should bring more new hair to cover their exposed eggs when a pied flycatcher is present near to tit nest than when a neutral (non-competing) species is present. We conducted decoy and playback experiment in Oulu and Turku, Finland. First, we removed and collected all the hair covering the tit eggs. Then, we measured how the perceived presence of flycatcher or waxwing (Bombycilla garrulus) affects tits' egg-covering by collecting and weighing the hair brought on the eggs and photographing the nest 24 h after the playback. RESULTS: Tits brought more hair into the nest and covered the eggs more carefully after flycatcher treatment, compared to waxwing treatment. We also found that the tits in Oulu (over 600 km to north from Turku) had more hair on the top of their eggs in general. CONCLUSIONS: Together, these results suggest that the counter-adaptation function of egg-covering against information parasites may be an extension of original function to protect eggs from low temperatures.

Observed fitness may affect niche overlap in competing species via selective social information use.

Social information transmission is important because it enables horizontal spread of behaviors, not only between conspecifics but also between individuals of different species. Because interspecific social information use is expected to take place among species with similar resource needs, it may have major consequences for the emergence of local adaptations, resource sharing, and community organization. Social information use is expected to be selective, but the conditions promoting it in an interspecific context are not well known. Here, we experimentally test whether pied flycatchers (Ficedula hypoleuca) use the clutch size of great tits (Parus major) in determining the quality of the observed individual and use it as a basis of decision making. We show that pied flycatchers copied or rejected a novel nest site feature preference of great tits experimentally manipulated to exhibit high or low fitness (clutch size), respectively. Our results demonstrate that the social transmission of behaviors across species can be highly selective in response to observed fitness, plausibly making the phenomenon adaptive. In contrast with the current theory of species coexistence, overlap between realized niches of species could dynamically increase or decrease depending on the observed success of surrounding individuals.

Flycatchers copy conspecifics in nest-site selection but neither personal experience nor frequency of tutors have an effect.

Using the behavior of others in guiding one's own behavior is a common strategy in animals. The prevailing theory predicts that young age and the inexperience of an individual are expected to increase the probability of adopting the behaviors of others. Also, the most common behavior in the population should be copied. Here, we tested the above predictions by examining social information use in the selection of nest-site features with a field experiment using a wild cavity nesting bird, the collared flycatcher (Ficedula albicollis). We used an experimental design in which geometric symbols depict nest-site features. By manipulating the apparent symbol choices of early settled individuals and monitoring the choices of later arriving birds, we can study social information use without bias from learned or innate preferences. Flycatchers were found to use social information in the selection of nest-site features, with about 60% of the population preferring the manipulated conspecific choices. However, age and experience as explanatory factors suggested by the social information use theory did not explain the choices. The present result, in concert with earlier similar experiments, implies that flycatchers may in some situations rely more on interspecific information in the selection of nest-site characteristics.

How does variation in the environment and individual cognition explain the existence of consistent behavioral differences?

According to recent studies on animal personalities, the level of behavioral plasticity, which can be viewed as the slope of the behavioral reaction norm, varies among individuals, populations, and species. Still, it is conceptually unclear how the interaction between environmental variation and variation in animal cognition affect the evolution of behavioral plasticity and expression of animal personalities. Here, we (1) use literature to review how environmental variation and individual variation in cognition explain population and individual level expression of behavioral plasticity and (2) draw together empirically yet nontested, conceptual framework to clarify how these factors affect the evolution and expression of individually consistent behavior in nature. The framework is based on simple principles: first, information acquisition requires cognition that is inherently costly to build and maintain. Second, individual differences in animal cognition affect the differences in behavioral flexibility, i.e. the variance around the mean of the behavioral reaction norm, which defines plasticity. Third, along the lines of the evolution of cognition, we predict that environments with moderate variation favor behavioral flexibility. This occurs since in those environments costs of cognition are covered by being able to recognize and use information effectively. Similarly, nonflexible, stereotypic behaviors may be favored in environments that are either invariable or highly variable, since in those environments cognition does not give any benefits to cover the costs or cognition is not able to keep up with environmental change, respectively. If behavioral plasticity develops in response to increasing environmental variability, plasticity should dominate in environments that are moderately variable, and expression of animal personalities and behavioral syndromes may differ between environments. We give suggestions how to test our hypothesis and propose improvements to current behavioral testing protocols in the field of animal personality.

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.