Do Fleas,and/or Old Nest Material,Influence Nest-site preference in Hole-nesting Passerines?

This study examined nest-site choice in a migratory population of pied flycatchers (Ficedula hypoleuca) and sympatric populations of three resident tit species (Parus major, P. caeruleus and P. palustris) in central Sweden. All four species are secondary-cavity nesters which naturally breed in pre-formed tree cavities but readily use artificial nest boxes. We asked whether flycatchers and tits discriminate between nest boxes that: 1. Are ‘empty’; 2. Contain old nests without ectoparasites (fleas Ceratophyllus sp.); or 3. Contain old nests with ectoparasites. We found that pied flycatchers preferred nest boxes containing old nests, regardless of whether these nests held parasites. In contrast, tits did not discriminate between the three types of boxes. Tits may pay a cost for their lack of choosiness: after the breeding season, tit nests contained more fleas than flycatcher nests. Nevertheless, parasites did not affect the choice of a nest site in any of the species studied. We suggest that the migratory flycatchers are in a hurry to start breeding upon arrival and use the presence of an old nest as a shortcut cue to assess nest-site quality. Also, they may save valuable time by copying the choice of previous breeders. Non-migratory tits may have more time to inspect nest sites, but do not seem to use the same cues in nest-site selection as the pied flycatcher.     

Are nest sites actively chosen? Testing a common assumption for three non-resource limited birds

Many widely-accepted ecological concepts are simplified assumptions about complex situations that remain largely untested. One example is the assumption that nest-building species choose nest sites actively when they are not resource limited. This assumption has seen little direct empirical testing: most studies on nest-site selection simply assume that sites are chosen actively (and seek explanations for such behaviour) without considering that sites may be selected randomly. We used 15 years of data from a nestbox scheme in the UK to test the assumption of active nest-site choice in three cavity-nesting bird species that differ in breeding and migratory strategy: blue tit (Cyanistes caeruleus), great tit (Parus major) and pied flycatcher (Ficedula hypoleuca). Nest-site selection was non-random (implying active nest-site choice) for blue and great tits, but not for pied flycatchers. We also considered the relative importance of year-specific and site-specific factors in determining occupation of nest sites. Site-specific factors were more important than year-specific factors for the tit species, while the reverse was true for pied flycatchers. Our results show that nest-site selection, in birds at least, is not always the result of active choice, such that choice should not be assumed automatically in studies of nesting behaviour. We use this example to highlight the need to test key ecological assumptions empirically, and the importance of doing so across taxa rather than for single “model” species.     

Climate change can alter competitive relationships between resident and migratory birds

Climate change could affect resource competition between resident and migratory bird species by changing the interval between their onsets of breeding or by altering their population densities. We studied interspecific nest-hole competition between resident great tits and migrant pied flycatchers in South-Western Finland over the past five decades (1953-2005). We found that appearance of fatal take-over trials, the cases where a pied flycatcher tried to take over a great tit nest but was killed by the tit, increased with a reduced interspecific laying date interval and with increasing densities of both tits and flycatchers. The probability of pied flycatchers taking over great tit nests increased with the density of pied flycatchers. Laying dates of the great tit and pied flycatcher are affected by the temperatures of different time periods, and divergent changes in these temperatures could consequently modify their competitive interactions. Densities are a result of reproductive success and survival, which can be affected by separate climatic factors in the resident great tit and trans-Saharan migrant pied flycatcher. On these bases we conclude that climate change has a great potential to alter the competitive balance between these two species.     

Ectoparasites,nest site choice and breeding success in the pied flycatcher

It has recently been suggested that nest box studies might bias the measurement of behavioural and life-history traits, because the removal of old nests may reduce the load of ectoparasites. This experimental artefact may have notable effects on nest site choice and breeding success in cavity-breeding birds. We tested (i) if pied flycatchers Ficedula hypoleuca prefer clean nest boxes and (ii) if old nest material affects the number of parasites and the breeding success of pied flycatchers. In the first experiment we offered birds one cleaned nest box and one nest box with old nest material from the previous year. The two nest boxes were placed in very similar sites near each other. In this experiment all pied flycatchers clearly preferred dirty nest boxes. In the second part of the study we distributed clean nest boxes and dirty ones on the same study area. After breeding was over we counted the number of fleas Ceratophyllus gallinae in the nest material. This flea species was the most abundant and probably the most hazardous parasite in the nests. Surprisingly, we found that there were significantly more fleas in the nest boxes with nests of the current year only than in the boxes with nests of both current and previous year. This might explain the preference for the dirty boxes. However, our results do not indicate that the number of fleas affects breeding success in the pied flycatcher.     

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca , at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus . After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.     

Antioxidants in eggs of great tits <Emphasis Type="Italic">Parus major</Emphasis> from Chernobyl and hatching success

Antioxidants are powerful protectors against the damaging effects of free radicals that constitute the inevitable by-products of aerobic metabolism. Growing embryos are particularly susceptible to the damaging effects of free radicals produced during rapid growth, and mothers of many species provide protection against such damage by allocating antioxidants to their eggs. Birds living in radioactively contaminated areas use dietary antioxidants to cope with the damaging effects of radiation, but females also allocate dietary antioxidants to eggs, potentially enforcing a physiological trade-off between self-maintenance and reproductive investment. Here we tested whether female great tits Parus major breeding in radioactively contaminated study areas near Chernobyl allocated less dietary antioxidants to eggs, and whether such reduced allocation of dietary antioxidants to eggs had fitness consequences. Concentrations of total yolk carotenoids and vitamins A and E were depressed near Chernobyl compared to concentrations in a less contaminated Ukrainian study area and a French control study area, and all antioxidants showed dose-dependent relationships with all three dietary antioxidants decreasing with increasing level of radiation at nest boxes. These effects held even when controlling statistically for potentially confounding habitat variables and covariation among antioxidants. Laying date was advanced and clutch size increased at nest boxes with high dose rates. Hatching success increased with increasing concentration of vitamin E, implying that hatching success decreased at boxes with high levels of radiation, eventually eliminating and even reversing the higher potential reproductive output associated with early reproduction and large clutch size. These findings are consistent with the hypothesis that radioactive contamination reduced levels of dietary antioxidants in yolks, with negative consequences for hatching success and reproductive success.     

Nest site preference depends on the relative density of conspecifics and heterospecifics in wild birds

Background

Social learning allows animals to eavesdrop on ecologically relevant knowledge of competitors in their environment. This is especially important when selecting a habitat if individuals have relatively little personal information on habitat quality. It is known that birds can use both conspecific and heterospecific information for social learning, but little is known about the relative importance of each information type. If provided with the choice between them, we expected that animals should copy the behaviour of conspecifics, as these confer the best information for that species. We tested this hypothesis in the field for Pied Flycatchers Ficedula hypoleuca arriving at their breeding grounds to select a nest box for breeding. We assigned arbitrary symbols to nest boxes of breeding pied flycatchers (conspecifics) and blue and great tits, Cyanistes caeruleus and Parus major (heterospecifics), in 2014 and 2016 in two areas with different densities of tits and flycatchers. After ca 50% of flycatchers had returned and a flycatcher symbol was assigned to their nest box, we gave the later arriving flycatchers the choice between empty nest boxes with either a conspecific (flycatcher) or a heterospecific (tit) symbol.

Results

As expected, Pied Flycatchers copied the perceived nest box choice of conspecifics, but only in areas that were dominated by flycatchers. Against our initial expectation, flycatchers copied the perceived choice of heterospecifics in the area heavily dominated by tits, even though conspecific minority information was present.

Conclusions

Our results confirm that the relative density of conspecifics and heterospecifics modulates the propensity to copy or reject novel behavioural traits. By contrasting conspecific and heterospecific ecology in the same study design we were able to draw more general conclusions about the role of fluctuating densities on social information use.

     

Positive feedback between ecological and reproductive character displacement in a young avian hybrid zone

Character displacement can reduce costly interspecific interactions between young species. We investigated the mechanisms behind divergence in three key traits-breeding habitat choice, timing of breeding, and plumage coloration-in Ficedula flycatchers. We found that male pied flycatchers became expelled from the preferred deciduous habitat into mixed forest as the superior competitor, collared flycatchers, increased in numbers. The peak in food abundance differs between habitats, and the spatial segregation was paralleled by an increased divergence in timing of breeding between the two species. Male pied flycatchers vary from brown to black with brown coloration being more frequent in sympatry with collared flycatchers, a pattern often proposed to result from selection against hybridization, that is, reinforcement. In contrast to this view, we show that brown male pied flycatchers more often hybridize than black males. Male pied flycatcher plumage coloration influenced the territory obtained in areas of co-occurrence with collared flycatchers, and brown male pied flycatchers experienced higher relative fitness than black males when faced with heterospecific competition. We suggest that allopatric divergence in resource defense ability causes a feedback loop at secondary contact where male pied flycatchers with the most divergent strategy compared to collared flycatchers are favored by selection.     

Arrival and onset of breeding of three passerine birds in eastern Finland tracks climatic variation and phenology of insects

Anthropogenic climate change poses a challenge to the annual cycles of migratory birds. It has become urgent to understand whether migratory birds are able to advance their spring phenology when the climate is warming and whether they are able to adjust these phenological phases to the spring phenology in their breeding areas. In this work, we studied long‐term trends in first arrival and onset of breeding for three passerine birds in eastern Finland; the pied flycatcher Ficedula hypoleuca, the common redstart Phoenicurus phoenicurus and the great tit Parus major. The pied flycatcher and the common redstart are long‐distance migrants while the great tit is a partial migrant in Finland. We asked what environmental variables best explain the first arrival or onset of breeding, if there is evidence of ‘thermal delay’ (long‐term increase in the accumulated temperatures) at arrival or onset of breeding and if the interannual variation in the onset of breeding correlates with variation in spring phenology of local insects. We found that the pied flycatcher and the common redstart had advanced their first arrival (explained by increased temperatures at the migration route), but we found no long‐term change in the onset of breeding (explained by local temperatures). Also, the onset of breeding of the great tit is tracking local temperatures. We found no or only weak evidence of thermal delay at arrival or onset of breeding for any of the species. The onsets of breeding for the pied flycatcher and the great tit are also closely tracking the spring phenology of the local insects. The stable or increasing population sizes of all three species in Finland could be a result from their ability to effectively track climatic and environmental variation.     

Effects of nocturnal illumination on life-history decisions and fitness in two wild songbird species

The effects of artificial night lighting on animal behaviour and fitness are largely unknown. Most studies report short-term consequences in locations that are also exposed to other anthropogenic disturbance. We know little about how the effects of nocturnal illumination vary with different light colour compositions. This is increasingly relevant as the use of LED lights becomes more common, and LED light colour composition can be easily adjusted. We experimentally illuminated previously dark natural habitat with white, green and red light, and measured the effects on life-history decisions and fitness in two free-living songbird species, the great tit (Parus major) and pied flycatcher (Ficedula hypoleuca) in two consecutive years. In 2013, but not in 2014, we found an effect of light treatment on lay date, and of the interaction of treatment and distance to the nearest lamp post on chick mass in great tits but not in pied flycatchers. We did not find an effect in either species of light treatment on breeding densities, clutch size, probability of brood failure, number of fledglings and adult survival. The finding that light colour may have differential effects opens up the possibility to mitigate negative ecological effects of nocturnal illumination by using different light spectra.     

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.     

Seasonal habitat-dependent change in nest box occupation by Tengmalm’s owl associated with a corresponding change in nest predation

We investigated the nest-site settlement of the forest-dwelling Tengmalm’s owl (Aegolius funereus) in the temperate zone of Central Europe (Czech Republic) over a 14-year period. We analysed the effect of changes in nest-site suitability (expressed as the proportion of deciduous forest) within the immediate surroundings of the nest boxes (a buffer radius of 25 m) in relation to nest box occupancy, the mean standardized laying date (further the laying date), occurrence of nest predation, and breeding density. In line with the predictions of the site dependence model, we found a positive relationship between the laying date and the proportion of deciduous trees in the nest box surrounding. Further, we found the interactive effect of the laying date and the proportion of deciduous forest surrounding a nest box on the occurrence of breeding attempts predated by pine martens. A higher proportion of deciduous forest was found in the surroundings of predated than non-predated breeding attempts, and a steeper positive relationship between the laying date and the proportion of deciduous forest was found in the surroundings of non-predated than predated breeding attempts. The change in the owls’ habitat utilization can be explained by the process of tree leafing which makes the deciduous forest a less suitable habitat with high risk of nest predation before tree foliage and a more suitable habitat with low risk of nest predation after tree foliage. We conclude that our results are in agreement with the site dependence model, and that the degree of foliage of deciduous forest in the nest surrounding of Tengmalm’s owl is a key factor determining site suitability due to predation risk from pine martens in Central Europe.     

Heterospecific attraction and food resources in migrants' breeding patch selection in northern boreal forest

We studied experimentally how heterospecific attraction may affect habitat selection of migrant passerine birds in Finnish Lapland. We manipulated the densities of resident tit species (Parus spp.). In four study plots residents were removed before the arrival of the migrants in the first study year, and in four other plots their densities were increased by releasing caught individuals. In the second year the treatments of the areas were reversed, allowing paired comparisons within each plot. We also investigated the relative abundance of arthropods in the study plots by the sweep-net method. This allowed us to estimate the effect of food resources on the abundance of birds. The heterospecific attraction hypothesis predicts that densities of migrant species (especially habitat generalists) would be higher during increased resident density. Results supported this prediction. Densities and number of the most abundant migrant species were significantly higher when resident density was increased than when they were removed. On the species level the redwing (Turdus iliacus) showed the strongest positive response to the increased abundance of tits. Migrant bird abundances seemed not to vary in parallel with relative arthropod abundance, with the exception of the pied flycatcher (Ficedula hypoleuca) which showed a strongly positive correlation with many arthropod groups. The results of the experiment indicate that migrants can use resident tit species as a cue to a profitable breeding patch. The relationship between the abundance of the birds and arthropods suggests that annual changes in food resources during the breeding season probably do not have a very important effect on bird populations in these areas. The results stress the importance of positive interspecific interactions in structuring northern breeding bird communities. Received: 1 September 1997 / Accepted: 22 January 1998     

Temporal differences in food abundance promote coexistence between two congeneric passerines

Many related species share the same environment and utilize similar resources. This is surprising because based on the principle of competitive exclusion one would predict that the superior competitor would drive the other species to extinction; coexistence is only predicted if interspecific competition is weaker than intraspecific competition. Interspecific competition is frequently reduced by differential resource use, resulting in habitat segregation. In this paper, we use the closely related collared and pied flycatcher to assess the potential of habitat differences to affect interspecific competition through a different mechanism, namely by generating temporal differences in availability of similar food resources between the two species. We found that the tree species composition of the breeding territories of the two species differed, mainly by a higher abundance of coniferous species around nest-boxes occupied by pied flycatchers. The temporal availability of caterpillars was measured using frass traps under four deciduous and two coniferous tree species. Deciduous tree species showed an early and narrow peak in abundance, which contrasted with the steady increase in caterpillar abundance in the coniferous tree species through the season. We subsequently calculated the predicted total caterpillar biomass available in each flycatcher territory. This differed between the species, with biomass decreasing more slowly in pied flycatcher territories. Caterpillar biomass is strongly correlated with the reproductive success of collared flycatchers, but much less so with pied flycatchers. However, caterpillar availability can only partly explain the differences in seasonal decline of reproductive success between the two species; we discuss additional factors that may contribute to this species difference. Overall, our results are consistent with the suggestion that minor habitat differences between these two species may contribute to promoting their coexistence.     

Does weather affect biting fly abundance in avian nests?

Environmental factors may strongly affect avian‐biting fly interactions in different ways because insects are heterothermic organisms that depend greatly on environmental variables to activate their metabolism and behaviour. We studied the effects of weather on both blackfly (Simuliidae) and biting midge Culicoides (Ceratopogonidae) abundance in nests of three passerine species: blue tits Cyanistes caeruleus, great tits Parus major and pied flycatchers Ficedula hypoleuca, breeding in the same area. We controlled for different host‐related factors (hatching date, brood size and host species). Blackfly abundance was negatively related to minimum temperature. In addition, blackfly and biting midge abundances were negatively affected by wind speed measured at 07.00 h, but blackfly abundance was positively associated to wind speed at 18.00 h. We found higher blackfly and biting midge abundances in nests with larger broods breeding later in the season, and significantly higher biting midge abundance in pied flycatcher nests as compared to tit nests. These results represent, to our knowledge, the first report of both environmental and host‐related effects on haematophagous fly abundance in the nests of wild hole‐nesting birds.     

Traditional versus non-traditional nest-site choice: alternative decision strategies for nest-site selection

In order to understand habitat selection, it is important to consider the way individual animals assess the suitability of a future reproductive site. One way of investigating mechanisms (such as those involved in nest site selection) is to examine breeding success and habitat characteristics in terms of animals returning to a place where they have already reproduced and using the same location over successive years or searching for new alternatives. This approach seems especially suitable for testing recent hypotheses suggesting that nest site selection is an integrative process that includes the use of social information (e.g. past breeding success of conspecifics). Determining the factors that elicit conservative or innovative behaviour regarding nest-site selection could be important for improving our understanding of habitat selection decisions in animals. More than half of the nests of the long-distance migratory lesser grey shrike Lanius minor, are built in the same or neighbouring trees. We found no evidence that habitat characteristics influence nest-site tradition. On the contrary, social information in terms of the presence of conspecifics and past reproductive success in terms of complete nest failures due to nest predation (but not detailed information such as variation in fledgling number) influenced nest-site tradition. Hence, social information and past reproductive success may play a role in nest-site choice in this species. Our results further demonstrate that previous experience with a nest site does not appear to be beneficial.     

Linking anti-predator behaviour and habitat quality: group effect in nest defence of a passerine bird

In habitats where the density of breeding individuals is higher, breeding success has been shown to increase with the number of close conspecific and heterospecific neighbours. However, the mechanisms linking habitat quality, group size of prey individuals and offspring defence are poorly known. In this field study, we examined the relationships between habitat quality and parental nest defence behaviour in the pied flycatcher (Ficedula hypoleuca). We found that mobbing is more intense in unmanaged forests where birds breed in more dense and diverse communities than in heavily managed young forests where heterospecific densities are lower. We also found that the mobbing activities of pied flycatchers breeding in unmanaged mature boreal forests attracted more neighbouring prey individuals than in nearby managed forests. This study shows that habitat quality-mediated effects might be responsible for the decreased group size of mobbing birds in managed forests, which may lead to less effective communal defence.     

Positive fitness consequences of interspecific interaction with a potential competitor

The coexistence of species sharing mutual resources is usually thought to be limited by negative processes such as interspecific competition. This is because an overlap in resource use leads to negative fitness consequences, and traits favouring avoidance of potential competitors, for example in habitat selection, are therefore selected for. However, species interactions are acknowledged to vary from negative (competition) to mutualism, although empirical evidence for positive interspecific interactions from natural communities of other than plants and sessile animals is scarce. Here, we experimentally examined the habitat selection and its fitness consequences of a migrant bird, the pied flycatcher (Ficedula hypoleuca), in relation to the presence of competitively superior birds, resident titmice (Parus spp.). Experiments were conducted on two spatial scales: landscape and nest-site scale. We demonstrate that pied flycatchers were attracted to and accrued fitness benefits from the presence of titmice. Flycatchers breeding in tight association with titmice initiated breeding earlier, had larger broods and heavier young than solitarily breeding flycatchers. This paradoxical result indicates that species interactions may switch from negative to positive and that the coexistence of species is not always restricted by negative costs caused by other species.     

Habitat selection responses of parents to offspring predation risk: an experimental test

The ability of nest predation to influence habitat settlement decisions in birds is widely debated, despite its importance in limiting fitness. Here, we experimentally manipulated nest predation risk across a landscape and asked the question, do migratory birds assess and respond to variation in nest predation risk when choosing breeding habitats? We examined habitat preference by quantifying the density and settlement date of eight species of migratory passerines breeding in areas with and without intact nest predator communities. We found consistently more individuals nesting in areas with reduced nest predation than in areas with intact predator assemblages, although predation risk had no influence on settlement or breeding phenology. Additionally, those individuals occupying safer nesting habitats exhibited increased singing activity. These findings support a causal relationship between habitat choice and nest predation risk and suggest the importance of nest predation risk in shaping avian community structure and breeding activity.     

Conspecific attraction and nest site selection in a nomadic species,the zebra finch

Conspecific nesting density affects many aspects of breeding biology, as well as habitat selection decisions. However, the large variations in breeding density observed in many species are yet to be fully explained. Here, we investigated the settlement patterns in a colonial species with variable breeding density and where resource distribution could be manipulated. The zebra finch, Taeniopygia guttata, is a classic avian model in evolutionary biology but we know surprisingly very little about nest site selection strategies and nesting densities in this species, and in fact, in nomadic species in general. Yet, important determinants of habitat selection strategies, including temporal predictability and breeding synchrony, are likely to be different in nomadic species than in the non‐nomadic species studied to date. Here, we manipulated the distribution of nesting sites (by providing nest boxes) and food patches (feeders) to test four non‐exclusive habitat selection hypotheses that could lead to nest aggregation: 1) attraction to resources, 2) attraction to breeding conspecifics, and 3) attraction to successful conspecifics and 4) use of private information (i.e. own reproductive success on a site). We found that wild zebra finches used conspecific presence and possibly reproductive success, to make decisions over where to locate their nests, but did not aggregate around water or food within the study areas. Moreover, there was a high degree of inter‐individual variation in nesting density preference. We discuss the significance of our results for habitat selection strategy in nomadic species and with respect to the differential selection pressures that individuals breeding at different densities may experience.