Sex allocation is color morph-specific and associated with fledging condition in a wild bird.

Melanin-based color polymorphism is predicted to evolve and maintain through differential fitness of morphs in different environments, and several empirical studies indicate that life history strategies, physiology, and behavior vary among color morphs. Sex allocation theory predicts that parents should adjust their sex allocation based on differential costs of raising sons and daughters, and therefore, color morphs are expected to modify their brood sex ratio decisions. In color polymorphic tawny owls (Strix aluco), the pheomelanistic brown morph is associated with higher energy requirements, faster growth, and higher parental effort than the gray morph. As hypothesized, we find that brown tawny owl mothers produced more daughters in early broods and more males in late broods, whereas gray mothers did the opposite. At fledging, daughters of early broods and of brown mothers were heavier than those of late broods or gray mothers. Hence, larger and more costly daughters appeared to benefit more than males from being born to brown mothers early in the season. Brown mothers breeding later in the season produced more cheap sons, while gray mothers face fewer challenges under limited resources and favor daughters. These findings suggest that environmental conditions influence brood sex allocation strategies of genetically determined color morphs differently.

Camouflage efficiency in a colour-polymorphic predator is dependent on environmental variation and snow presence in the wild.

Colour polymorphism can be maintained by colour morph-specific benefits across environmental conditions. Currently, the amount and the duration of snow cover during winter decrease especially in northern latitudes, which can alter the potential for camouflage of animals with light and dark morphs. Tawny owls, Strix aluco, are colour-polymorphic avian predators with dark (brown) and light (grey) colour morphs, where the grey morph is presumed to enjoy camouflage benefits under snowy conditions. We studied the camouflage potential of morphs in two tawny owls potential using passerines' probability to mob in the wild during winter with and without snow. For comparison with other seasons, we also repeated the experiment during spring and autumn. We found that grey tawny owls have a lower probability of being mobbed than the brown tawny owls only during snowy winters. The two colour morphs therefore experience differential benefits across snow conditions, which may help to maintain colour morphs in the population, although further warming of winter climate will reduce the potential for camouflage for grey tawny owls in northern latitudes.

Regional variation in climate change alters the range-wide distribution of colour polymorphism in a wild bird.

According to Gloger's rule, animal colouration is expected to be darker in wetter and warmer climates. Such environmental clines are predicted to occur in colour polymorphic species and to be shaped by selection if colour morphs represent adaptations to different environments. We studied if the distribution of the colour polymorphic tawny owl (Strix aluco) morphs (a pheomelanic brown and a pale grey) across Europe follow the predictions of Gloger's rule and if there is a temporal change in the geographical patterns corresponding to regional variations in climate change. We used data on tawny owl museum skin specimen collections. First, we investigated long-term spatiotemporal variation in the probability of observing the colour morphs in different climate zones. Second, we studied if the probability of observing the colour morphs was associated with general climatic conditions. Third, we studied if weather fluctuations prior to the finding year of an owl explain colour morph in each climate zone. The brown tawny owl morph was historically more common than the grey morph in every studied climate zone. Over time, the brown morph has become rarer in the temperate and Mediterranean zone, whereas it has first become rarer but then again more common in the boreal zone. Based on general climatic conditions, winter and summer temperatures were positively and negatively associated with the proportion of brown morph, respectively. Winter precipitation was negatively associated with the proportion of brown morph. The effects of 5-year means of weather on the probability to observe a brown morph differed between climate zones, indicating region-dependent effect of climate change and weather on tawny owl colouration. To conclude, tawny owl colouration does not explicitly follow Gloger's rule, implying a time and space-dependent complex system shaped by many factors. We provide novel insights into how the geographic distribution of pheomelanin-based colour polymorphism is changing.

Heritability and Parental Effects in Telomere Length in a Color Polymorphic Long-Lived Bird.

AbstractRelative telomere length (RTL), an indicator of senescence, has been shown to be heritable but can also be affected by environmental factors, such as parental effects. Investigating heritability as well as parental effects and rearing environment can help us to understand the factors affecting offspring telomeres. Moreover, how phenotypic parental traits linked with fitness can impact offspring RTL is still unclear. A phenotypic marker closely associated with physiological traits and fitness is melanin-based color polymorphism, which in tawny owl (Strix aluco) is highly heritable and strongly associated with adult telomere shortening and survival. We studied narrow-sense heritability (h2) of RTL, as well as the impact of parental age and color morph and their interaction on offspring RTL. Offspring RTL at fledging was strongly positively correlated with both mother RTL and father RTL at breeding. Offspring RTL was also negatively associated with father age, suggesting that older fathers sired offspring with shorter telomeres. Parental color morph did not explain offspring RTL, and there were no interactive effects of parental morph and age, despite previously documented morph-specific senescence patterns. Our results suggest that RTL is highly heritable and affected by paternal age but not related to color polymorphism. This suggests that either morph-specific telomere shortening as an adult does not result in significantly shorter telomeres in their gametes, or that parents compensate morph-specific senescence via parental care. Morph-specific patterns of telomere dynamics in polymorphic species may thus emerge from different life history strategies adopted in adulthood.

Cold winters have morph-specific effects on natal dispersal distance in a wild raptor.

Dispersal is a key process with crucial implications in spatial distribution, density, and genetic structure of species' populations. Dispersal strategies can vary according to both individual and environmental features, but putative phenotype-by-environment interactions have rarely been accounted for. Melanin-based color polymorphism is a phenotypic trait associated with specific behavioral and physiological profiles and is, therefore, a good candidate trait to study dispersal tactics in different environments. Here, using a 40 years dataset of a population of color polymorphic tawny owls (Strix aluco), we investigated natal dispersal distance of recruiting gray and pheomelanic reddish-brown (hereafter brown) color morphs in relation to post-fledging winter temperature and individual characteristics. Because morphs are differently sensitive to cold winters, we predicted that morphs' natal dispersal distances vary according to winter conditions. Winter temperature did not affect the proportion of brown (or gray) among recruits. We found that dispersal distances correlate with winter temperature in an opposite manner in the two morphs. Although the gray morph undertakes larger movements in harsher conditions, likely because it copes better with winter severity, the brown morph disperses shorter distances when winters are harsher. We discuss this morph-specific natal dispersal pattern in the context of competition for territories between morphs and in terms of costs and benefits of these alternative strategies. Our results stress the importance of considering the interaction between phenotype and environment to fully disentangle dispersal movement patterns and provide further evidence that climate affects the behavior and local distribution of this species.

Fledging Mass Is Color Morph Specific and Affects Local Recruitment in a Wild Bird.

AbstractEarly-life conditions may have long-lasting effects on life history. In color polymorphic species, morph-specific sensitivity to environmental conditions may lead to differential fitness. In tawny owls (Strix aluco), pheomelanin-based color polymorphism is expected to be maintained because the brown morph has higher adult fitness in warmer environments, while selection favors the gray morph under colder conditions. Here we investigate body mass at fledging and its consequences until adulthood in a population at the species' cold range margin. Using 40 years of data (1979-2017), we show that brown pairs, which mainly produce brown offspring consistent with a one-locus-two-alleles inheritance model, consistently raised heavier offspring than mixed (gray-brown) pairs and gray pairs. Offspring mass declined seasonally, except among offspring raised by brown pairs. Brown offspring could be heavier because of morph-specific parental care and/or offspring growth. Furthermore, mass at fledging is associated with fitness: the probability of local recruitment into the breeding population increased with higher mass at fledging, especially in mild winters and with favorable food conditions, although recruitment is not morph specific. Fledgling mass thus provides a fitness benefit in terms of recruitment probability that is modulated by environmental factors, which appear to level off any direct morph-specific recruitment benefits.

Climate change and perishable food hoards of an avian predator: Is the freezer still working?

Changing climate can modify predator-prey interactions and induce declines or local extinctions of species due to reductions in food availability. Species hoarding perishable food for overwinter survival, like predators, are predicted to be particularly susceptible to increasing temperatures. We analysed the influence of autumn and winter weather, and abundance of main prey (voles), on the food-hoarding behaviour of a generalist predator, the Eurasian pygmy owl (Glaucidium passerinum), across 16 years in Finland. Fewer freeze-thaw events in early autumn delayed the initiation of food hoarding. Pygmy owls consumed more hoarded food with more frequent freeze-thaw events and deeper snow cover in autumn and in winter, and lower precipitation in winter. In autumn, the rotting of food hoards increased with precipitation. Hoards already present in early autumn were much more likely to rot than the ones initiated in late autumn. Rotten food hoards were used more in years of low food abundance than in years of high food abundance. Having rotten food hoards in autumn resulted in a lower future recapture probability of female owls. These results indicate that pygmy owls might be partly able to adapt to climate change by delaying food hoarding, but changes in the snow cover, precipitation and frequency of freeze-thaw events might impair their foraging and ultimately decrease local overwinter survival. Long-term trends and future predictions, therefore, suggest that impacts of climate change on wintering food-hoarding species could be substantial, because their 'freezers' may no longer work properly. Altered usability and poorer quality of hoarded food may further modify the foraging needs of food-hoarding predators and thus their overall predation pressure on prey species. This raises concerns about the impacts of climate change on boreal food webs, in which ecological interactions have evolved under cold winter conditions.

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.

Maternal predator-exposure affects offspring size at birth but not telomere length in a live-bearing fish.

The perception of predation risk could affect prey phenotype both within and between generations (via parental effects). The response to predation risk could involve modifications in physiology, morphology, and behavior and can ultimately affect long-term fitness. Among the possible modifications mediated by the exposure to predation risk, telomere length could be a proxy for investigating the response to predation risk both within and between generations, as telomeres can be significantly affected by environmental stress. Maternal exposure to the perception of predation risk can affect a variety of offspring traits but the effect on offspring telomere length has never been experimentally tested. Using a live-bearing fish, the guppy (Poecilia reticulata), we tested if the perceived risk of predation could affect the telomere length of adult females directly and that of their offspring with a balanced experimental setup that allowed us to control for both maternal and paternal contribution. We exposed female guppies to the perception of predation risk during gestation using a combination of both visual and chemical cues and we then measured female telomere length after the exposure period. Maternal effects mediated by the exposure to predation risk were measured on offspring telomere length and body size at birth. Contrary to our predictions, we did not find a significant effect of predation-exposure neither on female nor on offspring telomere length, but females exposed to predation risk produced smaller offspring at birth. We discuss the possible explanations for our findings and advocate for further research on telomere dynamics in ectotherms.

Age and sex differences in numerical responses, dietary shifts, and total responses of a generalist predator to population dynamics of main prey.

Fluctuations in the abundance of main prey species might shape animal communities, by inducing numerical responses and dietary shifts in predators. Whether numerical responses and dietary shifts differ among individuals of different age and sex has so far gained little attention. These differences could affect how much predators consume main and alternative prey, thus causing variation in predation pressure on main and alternative prey species. We studied the effect of fluctuating main prey abundance (voles) in autumn on the age and sex composition of a food-hoarding population of Eurasian pygmy owls Glaucidium passerinum (327 individuals), and on the species composition of their food stores in western Finland during 2003-2017 (629 food stores). Numbers of yearlings (< 1-year old) of both sexes and adult (+ 1-year old) females increased with increasing vole abundance. During low vole abundance, adult owls stored more small birds and less small mammals than yearlings. Females stored more small mammals than males and showed a tendency to store less birds. The amount of consumed birds (the most important alternative prey), and in particular of crested, willow, great, and blue tits, increased with low vole densities. Our results show that numerical, functional, and total responses of pygmy owls, and probably also other vertebrate predators, to the availability of the main prey in winter are shaped by the age and sex composition of the predator population, which both show large spatio-temporal variation in boreal forests.

Maternal food supplementation and perceived predation risk modify egg composition and eggshell traits but not offspring condition.

Mothers may vary resource allocation to eggs and embryos, which may affect offspring fitness and prepare them for future environmental conditions. The effects of food availability and predation risk on reproduction have been extensively studied, yet their simultaneous impacts on reproductive investment and offspring early life conditions are still unclear. We experimentally manipulated these key environmental elements using a 2×2 full factorial design in wild, free-living pied flycatchers (Ficedula hypoleuca), and measured egg composition, eggshell traits and offspring condition. Eggs laid in food-supplemented nests had larger yolks and thicker shells independently of predation risk, while eggs laid in nests exposed to predator cues had lower levels of immunoglobulins, independent of food supplementation. In nests without predator cues, shell biliverdin content was higher in eggs laid in food-supplemented nests. Incubation was 1 day shorter in food-supplemented nests and shorter incubation periods were associated with higher hatching success, but there were no direct effects of maternal treatment on hatching success. To investigate the impact of maternal treatment (via egg composition) on the offspring, we performed full brood cross-fostering after hatching to unmanipulated nests. Maternal treatment did not significantly affect body mass and immunoglobulin levels of offspring. Our results suggest that although prenatal maternal cues affected egg composition, these egg-mediated effects may not have detectable consequences for offspring growth or immune capacity. Unpredictable environmental stressors may thus affect parental investment in the eggs, but parental care may level off costs and benefits of differential maternal egg allocation.

Hair cortisol concentration in Siberian flying squirrels is unrelated to landscape and social factors.

Intact ecosystems are being lost or modified worldwide, and many animal species are now forced to live in altered landscapes. A large amount of scientific studies have focused on understanding direct effects of habitat alterations on species occurrence, abundance, breeding success, and other life history aspects. Much less attention has been placed on understanding how habitat alterations impact on the physiology of species, e.g., via elevated chronic stress when living in an altered landscape. Here, we quantify the effects of individual age and sex, as well as effects of landscape and social factors on chronic stress of an endangered forest specialist species, the Siberian flying squirrel Pteromys volans. We collected hair samples over 2 years from across 192 flying squirrels and quantified their chronic stress response via cortisol concentrations. We then ran statistical models to relate cortisol concentrations with landscape and social factors. We show that cortisol concentrations in flying squirrels are neither affected by habitat amount and connectivity, nor by the density of conspecifics in the area. We however found that cortisol concentration was higher in adults than in pups, and in males compared with females. Lack of an effect of environmental factors on cortisol concentrations may indicate low physiological sensitivity to alterations in the surrounding environment, possibly due to low densities of predators that could induce stress in the study area. Further research should focus on possible effects of varying predator densities, alone and in interaction with landscape features, in shaping chronic stress of this and other species.

Antioxidant Enzyme Activities Vary with Predation Risk and Environmental Conditions in Free-Living Passerine Birds.

Prolonged physiological stress response may lead to an excessive production of reactive oxygen species (ROS) and ultimately to oxidative stress and severe fitness costs. We investigated whether natural variation in predation risk, induced by pygmy owls (Glaucidium passerinum), modifies the oxidative status of two free-living food-supplemented passerine bird species-the great tit (Parus major) and the willow tit (Poecile montanus)-in March 2012 and 2013. Predation risk significantly affected antioxidant enzyme activities of willow tits. Antioxidant enzyme activities (principal component factor 2 [PC2] representing glutathione-S-transferase and superoxide dismutase activities) were higher in high predation risk areas in 2013 than in low predation risk areas in the same year. Higher enzyme activities may suggest higher ROS production in birds living under high predation risk. In addition, antioxidant enzyme activities (PC2) were also higher in high predation risk areas in 2013 than in high predation risk areas in the previous year, 2012. This may represent variation in the risk represented by pygmy owls, which is probably inversely related to the natural fluctuations in the densities of their main prey, voles. In willow tits, PC1 (representing catalase, total glutathione, the ratio of reduced to oxidized glutathione, and protein carbonylation) was not affected by perceived predation risk, nor were antioxidant levels or enzyme activities in great tits. Higher enzyme activities observed in willow tits suggest that predator presence can modify the antioxidant status of avian prey, but the response also seem to be influenced by other environmental characteristics, like harsh winter conditions.

Species and abundance of ectoparasitic flies (Diptera) in pied flycatcher nests in Fennoscandia.

BACKGROUND: Birds host several ectoparasitic fly species with negative effects on nestling health and reproductive output, and with the capability of transmitting avian blood parasites. Information on the abundance and distribution of the ectoparasitic fly genera Ornithomya (Hippoboscidae) and Protocalliphora (Calliphoridae) in northern Europe is still generally poor, and we thus explored their geographic range and occurrence of these flies in the nests of a common avian model species, the pied flycatcher Ficedula hypoleuca. METHODS: Nests of F. hypoleuca were collected from 21 locations across Fennoscandia in summer 2013, across a latitudinal gradient (between 56 °N - 70 °N) and examined for the presence of fly puparia. Adult specimens of Ornithomya spp. were also collected for species identification. Fly species were identified morphologically and identifications confirmed with DNA barcoding. RESULTS: We found three species: two louse-flies - Ornithomya chloropus and O. avicularia - and one blow-fly, Protocalliphora azurea. The prevalence of O. avicularia was higher in southern latitudes and this species was not encountered beyond 62 °N whereas O. chloropus and P. azurea occurred across the whole range of latitudes. The prevalence of O. chloropus further increased with increasing distance from the coast - a pattern not documented before. The three fly species showed no interspecific associations in their prevalence. CONCLUSIONS: Our study revealed relatively high prevalence for all the species (O. chloropus 59 %, O. avicularia 20 %, P. azurea 32 %), and an interesting spatial pattern in the prevalence of the two louse fly species. Our sample did not indicate any major range shifts towards the north for the southern species as compared to the information from the past. Morphological identification of O. chloropus did not match with the corresponding sequences published in the GenBank and taxonomy of this group calls for further studies.

Large-scale geographical variation in eggshell metal and calcium content in a passerine bird (Ficedula hypoleuca).

Birds have been used as bioindicators of pollution, such as toxic metals. Levels of pollutants in eggs are especially interesting, as developing birds are more sensitive to detrimental effects of pollutants than adults. Only very few studies have monitored intraspecific, large-scale variation in metal pollution across a species' breeding range. We studied large-scale geographic variation in metal levels in the eggs of a small passerine, the pied flycatcher (Ficedula hypoleuca), sampled from 15 populations across Europe. We measured 10 eggshell elements (As, Cd, Cr, Cu, Ni, Pb, Zn, Se, Sr, and Ca) and several shell characteristics (mass, thickness, porosity, and color). We found significant variation among populations in eggshell metal levels for all metals except copper. Eggshell lead, zinc, and chromium levels decreased from central Europe to the north, in line with the gradient in pollution levels over Europe, thus suggesting that eggshell can be used as an indicator of pollution levels. Eggshell lead levels were also correlated with soil lead levels and pH. Most of the metals were not correlated with eggshell characteristics, with the exception of shell mass, or with breeding success, which may suggest that birds can cope well with the current background exposure levels across Europe.

Higher nest predation risk in association with a top predator: mesopredator attraction?

Breeding close to top predators is a widespread reproductive strategy. Breeding animals may gain indirect benefits if proximity to top predators results in a reduction of predation due to suppression of mesopredators. We tested if passerine birds gain protection from mesopredators by nesting within territories of a top predator, the Ural owl (Strix uralensis). We placed nest boxes for pied flycatchers (Ficedula hypoleuca) in Ural owl nest sites and in control sites (currently unoccupied by owls). The nest boxes were designed so that nest predation risk could be altered (experimentally increased) after flycatcher settlement; we considered predation rate as a proxy of mesopredator abundance. Overall, we found higher nest predation rates in treatment than in control sites. Flycatcher laying date did not differ between sites, but smaller clutches were laid in treatment sites compared to controls, suggesting a response to perceived predation risk. Relative nest predation rate varied between years, being higher in owl nest sites in 2 years but similar in another; this variation might be indirectly influenced by vole abundance. Proximity to Ural owl nests might represent a risky habitat for passerines. High predation rates within owl territories could be because small mesopredators that do not directly threaten owl nests are attracted to owl nest sites. This could be explained if some mesopredators use owl territories to gain protection from their own predators, or if top predators and mesopredators independently seek similar habitats.

Geographical variation in egg mass and egg content in a passerine bird.

Reproductive, phenotypic and life-history traits in many animal and plant taxa show geographic variation, indicating spatial variation in selection regimes. Maternal deposition to avian eggs, such as hormones, antibodies and antioxidants, critically affect development of the offspring, with long-lasting effects on the phenotype and fitness. Little is however known about large-scale geographical patterns of variation in maternal deposition to eggs. We studied geographical variation in egg components of a passerine bird, the pied flycatcher (Ficedula hypoleuca), by collecting samples from 16 populations and measuring egg and yolk mass, albumen lysozyme activity, yolk immunoglobulins, yolk androgens and yolk total carotenoids. We found significant variation among populations in most egg components, but ca. 90% of the variation was among individuals within populations. Population however explained 40% of the variation in carotenoid levels. In contrast to our hypothesis, we found geographical trends only in carotenoids, but not in any of the other egg components. Our results thus suggest high within-population variation and leave little scope for local adaptation and genetic differentiation in deposition of different egg components. The role of these maternally-derived resources in evolutionary change should be further investigated.

Geographical trends in the yolk carotenoid composition of the pied flycatcher (Ficedula hypoleuca).

Carotenoids in the egg yolks of birds are considered to be important antioxidants and immune stimulants during the rapid growth of embryos. Yolk carotenoid composition is strongly affected by the carotenoid composition of the female's diet at the time of egg formation. Spatial and temporal differences in carotenoid availability may thus be reflected in yolk concentrations. To assess whether yolk carotenoid concentrations or carotenoid profiles show any large-scale geographical trends or differences among habitats, we collected yolk samples from 16 European populations of the pied flycatcher, Ficedula hypoleuca. We found that the concentrations and proportions of lutein and some other xanthophylls in the egg yolks decreased from Central Europe northwards. The most southern population (which is also the one found at the highest altitude) also showed relatively low carotenoid levels. Concentrations of ß-carotene and zeaxanthin did not show any obvious geographical gradients. Egg yolks also contained proportionally more lutein and other xanthophylls in deciduous than in mixed or coniferous habitats. We suggest that latitudinal gradients in lutein and xanthophylls reflect the lower availability of lutein-rich food items in the northern F. hypoleuca populations and in montane southern populations, which start egg-laying earlier relative to tree phenology than the Central European populations. Similarly, among-habitat variation is likely to reflect the better availability of lutein-rich food in deciduous forests. Our study is the first to indicate that the concentration and profile of yolk carotenoids may show large-scale spatial variation among populations in different parts of the species' geographical range. Further studies are needed to test the fitness effects of this geographical variation.

Habitat selection as an antipredator behaviour in a multi-predator landscape: all enemies are not equal.

1. Breeding territory choice constitutes a crucial antipredator behaviour for animals that determines reproductive success and survival during the breeding season. On arrival to breeding grounds migrant prey face a multitude of 'waiting' predators already settled within the landscape. 2. We studied territory selection and reproductive investment of migrant pied flycatcher (Ficedula hypoleuca) relative to breeding pygmy owls (POs) (Glaucidium passerinum) and Tengmalm's owls (TOs) (Aegolius funereus). Diurnal POs present a greater predation threat to adult flycatchers (up to 80% songbirds in diet) compared with nocturnal TOs (up to 36%). 3. During territory selection, pied flycatchers strongly avoided POs (occupation: 42% in presence vs. 92% in absence of owl nest) but not TOs (80% vs. 75%). This suggests that flycatchers are able to distinguish between two potential predators, avoiding dangerous POs but not obviously responding to the less risky TOs. 4. Flycatchers responded to presence of PO nests with c. 4-day delay in the start of egg-laying. A significantly prolonged nest building period contributed to this potentially costly breeding delay. Flycatchers further significantly reduced initial reproductive investment in presence of POs by laying 8.2% smaller clutch sizes, even if laying date was controlled. No breeding delay and clutch size reduction was found relative to TO presence. 5. Our results highlight flexibility in breeding territory selection and reproductive strategies as antipredator responses to perceived risk in a multi-predator environment. This supports the idea that for prey, not all predators are equal.