Seasonal and urban effects on the endocrinology of a wild passerine

In order to maximize their fitness, organisms in seasonal environments rely on external cues to optimally time their life‐history stages. One of the most important zeitgeber to time reproduction is the photoperiod, but further environmental cues are assessed to fine‐tune reproduction due to year‐to‐year variation in environmental conditions. However, in urbanized environments, the pervasive artificial light at night has altered the natural signal of light and darkness. Accordingly, artificial light at night was repeatedly shown to affect avian reproductive physiology and to advance seasonal reproduction in birds. However, these experiments were mainly conducted in the absence of further environmental cues to facilitate the investigation of the mechanisms which are still poorly understood. Here, we investigate whether the endocrine system of free‐ranging European blackbirds (Turdus merula) correlates with the amount of artificial light at night along a rural to urban gradient while the birds still encounter complementary environmental cues including seasonal variation in day length and temperature. Testosterone and estrone were assessed as metabolites in fecal samples and corticosterone in blood from mist‐netted blackbirds. We demonstrate that seasonal fluctuations in abiotic factors, individual conditions, but also light at night affect the reproductive and stress physiology of wild European blackbirds. Elevated artificial night light intensities were significantly positively correlated with corticosterone and negatively with female estrone levels. No effects of artificial light were found for testosterone levels. Our results suggest that female blackbirds in particular perceive even low levels of artificial light at night as a weak but chronic stressor that interacts with the hypothalamic‐pituitary‐gonadal axis and leads to a reduced secretion of reproductive hormones. These findings point out that the impacts of light pollution are diverse and we only slowly disentangle its multiple effects on physiology, ecology, and biodiversity.     

Long-Term Effects of Chronic Light Pollution on Seasonal Functions of European Blackbirds (Turdus merula)

Light pollution is known to affect important biological functions of wild animals, including daily and annual cycles. However, knowledge about long-term effects of chronic exposure to artificial light at night is still very limited. Here we present data on reproductive physiology, molt and locomotor activity during two-year cycles of European blackbirds (Turdus merula) exposed to either dark nights or 0.3 lux at night. As expected, control birds kept under dark nights exhibited two regular testicular and testosterone cycles during the two-year experiment. Control urban birds developed testes faster than their control rural conspecifics. Conversely, while in the first year blackbirds exposed to light at night showed a normal but earlier gonadal cycle compared to control birds, during the second year the reproductive system did not develop at all: both testicular size and testosterone concentration were at baseline levels in all birds. In addition, molt sequence in light-treated birds was more irregular than in control birds in both years. Analysis of locomotor activity showed that birds were still synchronized to the underlying light-dark cycle. We suggest that the lack of reproductive activity and irregular molt progression were possibly the results of i) birds being stuck in a photorefractory state and/or ii) chronic stress. Our data show that chronic low intensities of light at night can dramatically affect the reproductive system. Future studies are needed in order to investigate if and how urban animals avoid such negative impact and to elucidate the physiological mechanisms behind these profound long-term effects of artificial light at night. Finally we call for collaboration between scientists and policy makers to limit the impact of light pollution on animals and ecosystems.     

Reproductive interactions of the Shiny Cowbird Molothrus bonariensis and the Yellow-hooded Blackbird Agelaius icterocephalus in Trinidad

The reproductive interactions of the Shiny Cowbird Molothrus bonariensis , a brood parasite, and the Yellow-hooded Blackbird Agelaius icterocephalus , a host of the cowbird, were studied In Trinidad, West Indies. We gathered information on the breeding biology of the Shiny Cowbird and the Yellow-hooded Blackbird, the frequency of use of the host species, and the effects of brood parasitism on host breeding success. Yellow-hooded Blackbirds are polygynous for the most part; males build nests and attempt to attract females to lay in them by means of song and visual displays directed towards the nests. This behaviour probably makes it easy for cowbirds to locate breeding birds and their nests. Cowbird eggs were found in 153 of 377 (40–3 %) blackbird nests located before the nestling stage. Shiny Cowbird parasitism of the Yellow-hooded Blackbird had little negative impact on host reproductive success, whereas predation accounted for the majority of nest failures. Vigilant nest defense by male blackbirds combined with colonial breeding apparently also minimized the extent of host egg damage and removal by cowbirds, and the parasitized and unparasitized nests were equally successful at producing blackbirds. Cowbirds most frequently parasitized the first or only nesting attempts in blackbird territories, and first or only nests were also successful more frequently than subsequent nests.     

Landscape features associated to wind farms increase mammalian predator abundance and ground-nest predation

Wind farm implementation is a rapidly growing source of landscape transformation that may alter ecological processes such as predator–prey interactions. We tested the hypothesis that wind farms increase the activity of nest predators and, ultimately, increment ground-nest predation rates. We placed 18 plots in Iberian shrub-steppes (11 at control and seven at wind farm sites), each one comprised nine artificial ground-nests (three quail eggs/nest). Artificial nests were placed during two events: at the beginning (April) and at the end (June) of the breeding season in 2016 (n?=?324 artificial nests). We estimated the relative abundance of avian and large mammalian predators in the surroundings of each plot and recorded nest fate after 12 days exposure. We also measured variables at landscape and microhabitat scale that potentially affect predator abundance and nest predation. Wind farm sites contained higher cover of gravel roads and more large mammalian predators. Moreover, the abundance of large mammalian predators increased with surrounding cover of both trees and gravel-roads. Avian predator abundance and nest predation rates did not differ between control and wind farm sites, though nest predation did increase with the surrounding cover of crops and gravel roads. Lastly, nest predation was higher at the end of the breeding season and decreased with moss and lichen cover. Our results support previous evidence on the increase of mammalian predator abundance as the surface area of gravel-roads increases, pointing towards a potential mechanism for wind farms leading to rise ground-nest predation. Future wind energy projects should minimize the development of gravel-roads for wind turbine access or maintenance.

     

Does light pollution alter daylength? A test using light loggers on free-ranging European blackbirds (Turdus merula)

Artificial light at night is one of the most apparent environmental changes accompanying anthropogenic habitat change. The global increase in light pollution poses new challenges to wild species, but we still have limited understanding of the temporal and spatial pattern of exposure to light at night. In particular, it has been suggested by several studies that animals exposed to light pollution, such as songbirds, perceive a longer daylength compared with conspecifics living in natural darker areas, but direct tests of such a hypothesis are still lacking. Here, we use a combination of light loggers deployed on individual European blackbirds, as well as automated radio-telemetry, to examine whether urban birds are exposed to a longer daylength than forest counterparts. We first used activity data from forest birds to determine the level of light intensity which defines the onset and offset of daily activity in rural areas. We then used this value as threshold to calculate the subjective perceived daylength of both forest and urban blackbirds. In March, when reproductive growth occurs, urban birds were exposed on average to a 49-min longer subjective perceived daylength than forest ones, which corresponds to a 19-day difference in photoperiod at this time of the year. In the field, urban blackbirds reached reproductive maturity 19 day earlier than rural birds, suggesting that light pollution could be responsible of most of the variation in reproductive timing found between urban and rural dwellers. We conclude that light at night is the most relevant change in ambient light affecting biological rhythms in avian urban-dwellers, most likely via a modification of the perceived photoperiod.     

Brood size manipulations in Great Horned Owls Bubo virginianus: are predators food limited at the peak of prey cycles?

The reproduction of raptors strongly depends on food resources. It is unclear whether predators experience superabundant food during cyclic peaks of prey populations. In order to test this hypothesis, four pairs of Great Horned Owls Bubo virginianus with two young were subjected to brood size manipulations during high densities of cyclic Snowshoe Hare Lepus americanus populations in southwestern Yukon, Canada. Broods older than 35 days were temporarily enlarged by one, and then by two, young. No effects were observed when one owlet was added, but the addition of two young resulted in significant weight losses in manipulated broods. Females with enlarged broods moved farther from their nest sites at night, presumably reflecting increased hunting effort, and also spent less time near the nest during the day. Food additions to enlarged broods returned the parental behaviour to normal. We conclude that these large predators did not experience superabundant food at this stage of the breeding season during a peak in cyclic prey.     

The Seasonal Dynamics of Artificial Nest Predation Rates along Edges in a Mosaic Managed Reedbed

Boundaries between different habitats can be responsible for changes in species interactions, including modified rates of encounter between predators and prey. Such ‘edge effects’ have been reported in nesting birds, where nest predation rates can be increased at habitat edges. The literature concerning edge effects on nest predation rates reveals a wide variation in results, even within single habitats, suggesting edge effects are not fixed, but dynamic throughout space and time. This study demonstrates the importance of considering dynamic mechanisms underlying edge effects and their relevance when undertaking habitat management. In reedbed habitats, management in the form of mosaic winter reed cutting can create extensive edges which change rapidly with reed regrowth during spring. We investigate the seasonal dynamics of reedbed edges using an artificial nest experiment based on the breeding biology of a reedbed specialist. We first demonstrate that nest predation decreases with increasing distance from the edge of cut reed blocks, suggesting edge effects have a pivotal role in this system. Using repeats throughout the breeding season we then confirm that nest predation rates are temporally dynamic and decline with the regrowth of reed. However, effects of edges on nest predation were consistent throughout the season. These results are of practical importance when considering appropriate habitat management, suggesting that reed cutting may heighten nest predation, especially before new growth matures. They also contribute directly to an overall understanding of the dynamic processes underlying edge effects and their potential role as drivers of time-dependent habitat use.     

Sleepless in Town – Drivers of the Temporal Shift in Dawn Song in Urban European Blackbirds

Organisms living in urban environments are exposed to different environmental conditions compared to their rural conspecifics. Especially anthropogenic noise and artificial night light are closely linked to urbanization and pose new challenges to urban species. Songbirds are particularly affected by these factors, because they rely on the spread of acoustic information and adjust their behaviour to the rhythm of night and day, e.g. time their dawn song according to changing light intensities. Our aim was to clarify the specific contributions of artificial night light and traffic noise on the timing of dawn song of urban European Blackbirds (Turdus merula). We investigated the onset of blackbird dawn song along a steep urban gradient ranging from an urban forest to the city centre of Leipzig, Germany. This gradient of anthropogenic noise and artificial night light was reflected in the timing of dawn song. In the city centre, blackbirds started their dawn song up to 5 hours earlier compared to those in semi-natural habitats. We found traffic noise to be the driving factor of the shift of dawn song into true night, although it was not completely separable from the effects of ambient night light. We additionally included meteorological conditions into the analysis and found an effect on the song onset. Cloudy and cold weather delayed the onset, but cloud cover was assumed to reflect night light emissions, thus, amplified sky luminance and increased the effect of artificial night light. Beside these temporal effects, we also found differences in the spatial autocorrelation of dawn song onset showing a much higher variability in noisy city areas than in rural parks and forests. These findings indicate that urban hazards such as ambient noise and light pollution show a manifold interference with naturally evolved cycles and have significant effects on the activity patterns of urban blackbirds.     

Implications of nest-site limitation on density-dependent nest predation at variable spatial scales in a cavity-nesting bird

Nest‐site limitation may have different implications in the spatial distribution of breeding pairs depending on the availability of suitable habitat and the types of nest‐sites. Distribution of cavities suitable as nest sites may allow circumstantial aggregation or active choice of colonial nesting, which may have different implications on breeding performance through effects on breeding density, with variable costs and benefits depending on the consequences of intraspecific competition, social interactions and predation. We evaluated the effects of breeding density derived from nesting site limitation on breeding performance and predation at different spatial scales and considering multiple social, population and environmental limiting factors in the red‐billed chough Pyrrhocorax pyrrhocorax. The results indicate that variable breeding density may arise within the population depending on the availability and spatial distribution of nest‐sites. Nest‐site availability and distribution may also determine social breeding systems (isolated or aggregated) at variable densities, thus resembling differences found at different spatially distant populations under contrasting environmental conditions. Breeding performance was related to density‐dependent processes of population regulation, especially density‐dependent nest predation due to predator attraction to nest clusters. Results also indicate that predation pressure depend on density patterns at large scales. This suggest that predation may have important consequences on population dynamics of spatially structured populations depending on the strength of this kind of density dependence, which in turn may depend on habitat features affecting the prey but also the spatially variable guild of predators. Because habitat and nesting site availability may vary spatially depending on multiple human influences, understanding the strength and form in which breeding density and nest predation at different spatial scales may influence the size and persistence of populations can help to manage them more adequately.     

Dark tree cavities – a challenge for hole nesting birds?

Holes provide the safest nest sites for birds, but they are an underutilized resource; in natural forests there are usually more holes than birds that could use them. Some bird species could be prevented from nesting in holes because of their inability to operate in the low light conditions which occur in cavities. As no visual system can operate in complete darkness some nest cavities could be too dark to be useable even by hole‐nesters. Thus, the light conditions within tree cavities could constrain both the evolution of the hole nesting habit, and the nest site choice of the hole‐nesting birds. These ideas cannot be tested because little is known about the light conditions in cavities. We took an opportunity provided by ongoing studies of marsh tits Poecile palustris and great tits Parus major breeding in a primeval forest (Bia?owie?a National Park, Poland) to measure illumination inside their nest cavities. We measured illuminance in cavities at daybreak, which is just after the parents commenced feeding nestlings. Only ca 1% of incoming light reached the level of the nest. Illuminance at nests of both species (median = 0.1–0.2 lx) fell within mesopic‐scotopic range, where colour vision is impaired. Measurements in model cavities showed strong declines in illumination with distance from the entrance, with light levels typically as low as 0.01 lx at 40 cm from the cavity entrance. Thus cavities can be very dark, often too dark for the use of colour vision, and we suggest that ‘lighting’ requirements can affect the adoption of specific nest sites by hole nesting birds. We discuss implications of the findings for understanding the adaptations for hole‐breeding in birds.     

Independent trophic behavior and breeding success of a resident flycatcher and a coexisting migratory congener

Historical biases towards the study of Palearctic and Nearctic bird migration systems has resulted in a dearth of information on the ecology of intratropical migrants, which likely also play important ecological roles within their communities. For instance, there is little information on the foraging ecology and breeding biology of co‐occurring migrants and residents within the intratropical migratory system. Thus, we used two congeneric flycatchers – the resident Plain‐crested Elaenia Elaenia cristata (Aves: Tyrannidae), and the migrant Lesser Elaenia E. chiriquensis (Tyrannidae) – to check whether either of them showed foraging niche plasticity mediating their coexistence and the effects of such syntopy on their breeding behavior and fitness. To do so, we used an ensemble of methods that included the evaluation of food resource phenology, foraging behavior observations, stable isotope ratio analysis in birds’ tissues, and nest monitoring. Our results confirmed that residents have foraging behavior plasticity but reveals its trophic behavior and breeding success is independent of the coexistence with migrants. Since such results depart from the predictions based on the current (and often Northern‐biased) theories of bird migration, we discuss alternative hypotheses explaining our findings, including the influence of physiology on diet and the behavioral responses of nest predators. Once there is a clear knowledge gap about the interspecific interactions between residents and intratropical migrants, our study represents a simple, yet important, step towards understanding the mechanisms underlying this system.     

Nest predation and habitat change interact to influence Siberian jay numbers

We studied Siberian jays, breeding in northern Sweden, to examine the potential for interactions between nest predation and reduced vegetation heterogeneity around nest sites to cause a decrease in jay numbers. Parent behaviour and nests are highly cryptic in the species. Our 12-year data showed, however, that nests had a probability of only 0.46 to be successful and produce at least one nestling. Nest predation was intense and a main cause of nest failure. All predators that could be identified were visually oriented hunters, mostly other corvids able to colonize taiga forest only close to human settlements. Consistent with the idea that predators used visual cues, nest predation increased with parental activity, which suggests that predators used parental provisioning trips to locate nests. Furthermore, a reduction in daily nest survival rates with decreasing amount of nesting cover was more pronounced in areas with high corvid activity as predicted when cover mediates the hunting efficiency of visual oriented predators. Declining temperatures interacted with the effects of habitat characteristics to further reduce daily nest survival rates suggesting that parents were not able to increase nest visitation rates to satisfy the higher energy demands of their nestlings without endangering the nest. Our results identify a mechanism through which predation and human-induced reduction in nesting cover on a larger scale may interact to cause a reduction in Siberian jay numbers larger than expected from habitat loss alone.     

Nest height is affected by lamppost lighting proximity in addition to nestbox size in urban great tits

Both natural and artificial light have proximate influences on many aspects of avian biology, physiology and behaviour. To date artificial light at night is mostly considered as being a nuisance disrupting for instance sleep and reproduction of diurnal species. Here, we investigate if lamppost night lighting affects cavity‐nesting bird species inside their breeding cavity. Nest height in secondary cavity‐nesting species is the result of trade‐offs between several selective forces. Predation is the prevailing force leading birds to build thin nests to increase the distance towards the entrance hole. A thin nest may also limit artificial light exposure at night. Yet, a minimum level of daylight inside nesting cavities is necessary for adequate visual communication and/or offspring development. Against this background, we hypothesised that avian nest‐building behaviour varies in response to a change in night lighting. We monitored nest height of urban great tits Parus major during six years and found that it varied with artificial light proximity. The birds built thinner nests inside nestboxes of various sizes in response to increasing lamppost night light availability at the nest. In large nestboxes, the nests were also thinner when a lamppost was present in the territory. Whether this relationship between artificial night lighting and nest height reflects a positive or negative effect of urbanisation is discussed in the light of recent experimental studies conducted in rural populations by other research groups.     

Brood Value and Life Expectancy as Determinants of Parental Investment in Male Three-spined Sticklebacks,Gasterosteus aculeatus

29 breeding male three-spined sticklebacks (Gasterosteus aculeatus) caught in the wild at different times in the breeding season were exposed simultaneously to a potential threat to their nest (a conspecific male) and to two hunting trout (potential predators of adult sticklebacks, seen through a transparent partition). By promoting a variety of protective responses, the presence of the predators reduced both the time spent confronting the intruder and the rate at which it was attacked. Subjects with a clutch of eggs in their nest maintained higher levels of territorial defence in the presence of predators than did those with an empty nest. However, those breeding later in the season took fewer risks to defend their nest. Possible proximate mechanisms responsible for these results are discussed. In addition, the adaptive significance of the behavioural changes is considered in the light of the value of the brood and the expected future reproductive output of the sticklebacks breeding at different times in the season.     

Provision of supplementary food for wild birds may increase the risk of local nest predation

In countries such as the UK, USA and Australia, approximately half of all households provide supplementary food for wild birds, making this the public's most common form of active engagement with nature. Year‐round supplementary feeding is currently encouraged by major conservation charities in the UK as it is thought to be of benefit to bird conservation. However, little is understood about how the provision of supplementary food affects the behaviour and ecology of target and non‐target species. Given the scale of supplementary feeding, any negative effects may have important implications for conservation. Potential nest predators are abundant in urban areas and some species frequently visit supplementary feeding stations. We assess whether providing supplementary food affects the likelihood of nest predation in the vicinity of the feeder, by acting as a point attractant for potential nest predators. We provided feeding stations (empty, peanut feeder, peanut feeder with guard to exclude potential nest predators) in an area of suburban parkland in the UK and monitored the predation rate of eggs placed in artificial nests located at distances that replicated the size of typical suburban gardens. Nest predators (Magpies Pica pica, Grey Squirrels Sciurus carolinensis) were frequent visitors to filled feeders, and predation caused by Magpies, European Jays Garrulus glandarius and Grey Squirrels was significantly higher when nests were adjacent to filled feeders. The presence of a feeder guard did not significantly reduce nest predation. As supplementary feeding is becoming increasingly common during the breeding season in suburban habitats, we suggest that providing point attractants to nest predators at this time may have previously unconsidered consequences for the breeding success of urban birds.     

Light data from geolocation reveal patterns of nest visit frequency and suitable conditions for efficient nest site monitoring in Common Swifts Apus apus

ABSTRACT

Capsule: We developed a protocol for efficient monitoring of potential Common Swift Apus apus nest sites which considers variation in nest visit frequency across the breeding season and in relation to time of day and weather.

Aims: To investigate patterns of nest visit frequency in Common Swifts in order to improve the efficiency and reliability of the monitoring of nest sites threatened by building renovations.

Methods: We derived information on nest attendance from light data recorded by geolocators from ten adult Common Swifts during three breeding seasons (n?=?686 individual sampling days) and analysed how nest visit frequency varied across the breeding season and in relation to time of day and weather.

Results: The mean nest visit frequency was 5.63 visits per bird per day (0.32 visits per hour of daylight). The daily number of visits was highest at the beginning of July during chick-rearing. Moreover, it was positively correlated with temperature and negatively correlated with rainfall and wind speed. Nest visit frequency showed a distinct peak around sunset, while also being relatively high in the morning and around noon.

Conclusion: We recommend monitoring potential Common Swift nest sites in Central Europe between the end of June and mid-July during good weather between 0.50 and 7.75?h after sunrise or between 3.00?h before sunset and sunset, when observation bouts of 0.5–2.0?h provide an encounter probability greater than 90%. Our study shows that repurposing geolocator light data – usually used to study bird migration – for investigating nest attendance in cavity-breeding birds can provide important information for bird conservation.     

Female-biased incubation and strong diel sex-roles in the Two-banded Plover Charadrius falklandicus

The relative contributions of males and females to incubation, and the diel schedules by which incubation is shared, are important breeding system traits. We used infra-red sensitive cameras to record incubation patterns at 13 nests of the Two-banded Plover Charadrius falklandicus in the Falkland Islands during both day and night. Because predation risk can affect incubation behaviour, we also recorded the diel pattern of nest predation in the wider study population. We found high nest attendance, female-biased incubation, and strong diel sex-roles, with females incubating during the day and males at night. We also found that incubation intermissions tended to be short but frequent, and were correlated strongly with the diel pattern of nest predations which occurred exclusively in the daylight hours (probably due to the absence of terrestrial mammals from the study site). Our results suggest that sex-roles are unusually strict in the Two-banded Plover, and that these strict sex-roles lead to inequality in incubation sharing and the level of exposure to sources of energetic cost such as disturbance by nest predators.     

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.     

An effective nest trap for female Yellow-headed Blackbirds

ABSTRACT.    Capturing breeding female blackbirds using mist nets and food-baited traps can be challenging because nets are difficult to set in wetland habitats and females feed preferentially on insects. Given the problems associated with other trapping methods, we designed a highly effective nest trap that allowed us to successfully capture targeted female Yellow-headed Blackbirds ( Xanthocephalus xanthocephalus ) at our wetland study sites in North Dakota. From 15 May to 16 July 2004, we captured 56 of 59 (95%) targeted females during incubation and eight of 18 (44%) targeted females with nestlings, and were also able to recapture 12 of 16 females (75%). Based on these results, we recommend using our trap to capture female blackbirds during incubation rather than during the nestling period. The nest trap is easy and inexpensive to construct and is light and easy to carry through dense wetland vegetation. In addition, the trap's large diameter and wire construction provide the flexibility needed to trap at sites where the density of vegetation varies. This trap has also been used to capture female Red-winged Blackbirds ( Agelaius phoeniceus ) and could potentially be used to trap females of other species of wetland-breeding icterids, including Common Grackles ( Quiscalus quiscula ).     

Natural cavity restoration as an alternative to nest box supplementation

Nest box supplementation is widely used to increase nest‐site availability for cavity nesting animals but the analysis of its effects on individuals breeding in natural cavities is often neglected. This study offers a novel restoration technique to revert abandonment of natural breeding sites by a secondary cavity avian bird, the European roller (Coracias garrulus), and other ecologically similar species. We found that, after a program of nest box supplementation with ensuing monitoring, rollers gradually abandon nesting in natural and seminatural cavities in favor of nest boxes because the latter are of higher quality. We examine whether reducing the entrance size of natural and seminatural cavities improves their suitability for rollers. A 6‐year program reduced the diameter of the entrance of sandstone cavities and cavities in bridges. This led to a high occupancy (59%) of manipulated nest‐sites. Manipulated sites were most frequently occupied by rollers and little owls (Athene noctua) (31 and 18% of sites, respectively). Manipulation did not affect clutch size or fledgling success. We suggest that nest‐site diversity and nesting in natural cavities should be preserved to reduce nest box dependence. Our study illustrates the value of nest boxes when used alongside restoration of natural breeding sites and provides insights for the management of natural cavities.