Breeding success and lutein availability in great tit (Parus major)

The relationship among temporal variation in the availability of carotenoid-rich food, tissue carotenoid levels and breeding success are poorly known. We studied how diet quality and quantity affect the carotenoid profile and fledging success of great tit (Parus major) nestlings along a pollution gradient. We found declining seasonal trend in lutein concentration of caterpillars, which may be the explanation for the declining trend in nestlings' lutein concentration of plasma with season, despite the increase in caterpillar biomass. This may be because the biomass of most lutein-rich caterpillars (autumnal moths) decreased and less lutein-rich caterpillars (sawflies) increased during the breeding season. The temporal difference in occurrence of different caterpillar species means that peak lutein availability does not coincide with peak caterpillar abundance. However the positive association between total larval biomass and the number of great tit fledglings may suggest that fledging success depends more on total caterpillar availability than on lutein concentration of caterpillars.     

Habitat quality, nestling diet, and provisioning behaviour in great tits Parus major

Variation in early nutrition is often a strong predictor of offspring condition and fitness. In the case of woodland passerine birds, nestling diet is determined by the spatiotemporal distribution of prey items such as caterpillars during the nestling period, and is usually quantified as differences in provisioning behaviour between habitats. However, the habitat level does not account for variation between individual territories, the level at which competition and selection are assumed to operate. Here we use nestbox cameras and Radio Frequency Identification technology (RFID) to simultaneously assess variation in both nestling diet (components) and provisioning rates (quantity) among a sample (n=22) of different quality great tit Parus major territories selected from a larger breeding population (n=310 fledged broods) in a single year. Caterpillars were by far the most numerous item provisioned to nestlings (mean=75% of prey items), as expected given the known importance of this food source for this species. Broods raised close to an oak tree, or far from the woodland edge, were provisioned the highest proportion of caterpillars. Provisioning rates declined seasonally and there was a weak association between low provisioning rates and caterpillar rich diets. During the first week of the nestling stage, nestling condition was unrelated to the proportion of caterpillars in the diet, provisioning rates and oak proximity. Condition at fledging was slightly improved in broods fed a higher proportion of caterpillars in the diet and in broods raised close to an oak tree. However, in our data early breeding was the only predictor of recruitment success, although power was low for this test. Analyses of long‐term data (41 years) from the same population confirmed a relationship between oak proximity and fledgling mass, but not recruitment success. Our results suggest that territory level environmental variation can affect offspring condition, probably through observed changes in nestling diet, but that such variation does not necessarily produce discernable effects on offspring fitness.     

Age-Related Changes in Nestling Diet of the Cooperatively Breeding Green Woodhoopoe

In many socially monogamous bird species, parents of altricial young respond to the increasing demands of growing nestlings by increasing their feeding rate and the size of prey items delivered and by altering the types of prey provided. In some cooperatively breeding species, similar changes in feeding rate and prey size have been documented. However, potential changes in the types of prey delivered, both as nestlings age and by different group members, remain largely unexplored. Moreover, studies rarely compare the diet fed to nestlings with that eaten by the provisioning adults themselves. Here, I show that green woodhoopoe ( Phoeniculus purpureus ) nestlings receive a smaller proportion of spiders and larger proportions of caterpillars and centipedes as they grow older. Both male and female adults delivered a higher proportion of spiders to young nestlings than they ate while self-feeding, probably in response to particular nutritional requirements of the chicks. However, only males altered the proportions of caterpillars and centipedes delivered, providing smaller proportions to young nestlings than eaten themselves. These prey items may be too large for young nestlings to handle, and males may make a greater adjustment in provisioning diet than females because they collect more caterpillars and centipedes than do females. Although there were sex differences in provisioning diet, there were no differences between same-sex breeders and helpers in terms of the overall proportions of prey delivered or the changes with nestling age. Hence, individuals of different reproductive status may be following the same provisioning rules, at least in terms of prey type.     

Are colorful males of great tits Parus major better parents? Parental investment is a matter of quality

Given the known influence of parental investment on breeding success of great tits Parus major, females should be expected to use male parental quality as an essential criterion in mate choice. Since parental quality cannot usually be observed directly at the time of pairing, it has been suggested that females rely on male ornaments as indicative of their ability to provide parental care. This hypothesis, called the good parent hypothesis, has been tested repeatedly assessing only parental effort as the number of feedings made by parents. However, in evaluating parental investment, the focus should also be on the quality of prey captured rather than only on its quantity. We analyzed feeding rates and the provisioning of different prey in relation to both male yellow carotenoid-based breast coloration and the size of the black melanin-based stripe in a Mediterranean great tit population. We predicted that more carotenoid ornamented individuals would feed nestlings with a diet consisting of a higher proportion of caterpillars. However, and contrary to predictions, we found that males with higher values of hue in the yellow breast feathers, fed their offspring with a lower proportion of caterpillars and a higher proportion of spiders. In addition, nestlings that received a higher proportion of spiders showed an improved body condition after controlling for tarsus length and other variables. Male feeding rates correlated positively with brood size and tended to correlate negatively with date, although we did not find any effect of male coloration. Our data therefore support the good parent hypothesis, insofar as parental investment is also a matter of quality, and that, at least in the Mediterranean area, caterpillars are not the only key food source.     

Importance of feeding ecology to the reproductive success of Blackbirds Turdus merula nesting in rural habitats

The feeding ecology of Blackbirds Turdus merula breeding in contiguous woodland and farmland habitats was studied over three years. The aim of the study was to investigate how reproductive success was influenced by nestling diet and the provisioning rates of parents feeding nestlings. Parental provisioning rates increased with brood size, and consequently individual nestlings were no lighter in larger broods. None of the environmental factors measured had strong effects on parental provisioning rate. The nestling diet was dominated by caterpillars and earthworms, the former occurring in a short period in the middle of the breeding season. The availability of earthworms was higher in woodland and was dependent on rainfall in farmland. Nestling mass and provisioning rates were marginally higher under predominantly earthworm diets. Nestling mass increased with rainfall in farmland only, and was higher in farmland than in woodland or woodland-edge, although it is doubtful whether this result is of any significance for fledgling survival. Overall, Blackbirds were able to provision their nestlings adequately throughout the breeding season across a range of conditions. There was no evidence to suggest that reproductive success was constrained by aspects of feeding ecology within the natural range of brood size.     

Azure‐winged Magpies Cyanopica cyanus trade off reproductive success and parental care by establishing a size hierarchy among nestlings

It is common in birds that the sizes of nestlings vary greatly when multiple young are produced in one nest. However, the methods used by parents to establish size hierarchy among nestlings and their effect on parental provisioning pattern may differ between species. In the Azure‐winged Magpie Cyanopica cyanus, we explored how and why parents controlled the sizes of nestlings. Asynchronous hatching was the main cause of size hierarchy within the brood, although the laying of larger eggs later in the laying sequence reduced this effect. Parents with asynchronous broods produced more eggs and fledged more nestlings than those with synchronous broods but their brood provisioning rates, food delivery per feeding bout and feeding efficiency did not differ. We performed a cross‐fostering experiment to synchronize some asynchronous broods. Provisioning rates of asynchronous broods were lower than those of synchronized broods, but the daily growth rates and fledging body mass of their nestlings were not different. Our findings indicate that parents of asynchronous broods can achieve higher reproductive success than those of synchronous broods based on the same parental care, and the same reproductive success as those of synchronized broods based on less parental care. It appears that parent birds can better trade off reproductive success and parental care by establishing a size hierarchy among nestlings.     

Habitat fragmentation influences nestling growth in Mediterranean blue and great tits

In patchy forest areas, the size of the forest patch where birds breed has a strong influence on their breeding success. However, the proximate effects contributing to lowering the breeding success in small forest patches remain unclear; and a shortage of crucial resources in those forest patches has been suggested to account in some degree for this failure. With the aim to further investigate this issue, we have monitored the breeding cycle of blue and great tits in three ‘large’ forest patches (ranging between 26.5 and 29.6 ha) and twelve ‘small’ forest patches (ranging between 1.1 and 2.1 ha) in a Mediterranean area in central Spain, during three years (2011–2013). We also recorded the nestling diet inside the nest-boxes with the aid of handy-cams. Only males significantly differed between forest patch size categories; being on average younger and with better body condition in small patches for great and blue tits respectively. Reproductive traits did not vary between forest patch size categories, but the body condition of blue tit nestlings and the size of great tit nestlings did, being significantly better and larger respectively in large forest patches. The recruitment rate of blue tit nestlings was also higher in large patches. Regarding nestling diet, blue tits did not differ but great tits did, delivering a larger amount of caterpillars in large forest patches. Most variation in the reproductive traits occurred between years, probably due to annual differences in environmental conditions. This study suggests that food supply could be limiting the breeding success of birds above all in small patches, but also in large patches under particular environmental conditions.     

Brood provisioning rates and fledgling behavior of Cordilleran Flycatchers in southwestern Colorado

The behavior of young songbirds after fledging is one of the least understood phases of the breeding cycle, although parental provisioning rates and movement of fledglings are key to understanding life history evolution. We studied Cordilleran Flycatchers (Empidonax occidentalis) at two sites in southwestern Colorado, USA, from 2012 to 2017. We banded and sexed breeding adults to determine the relative contributions of males and females to nestling and fledgling care, and attached radio‐transmitters to nestlings to facilitate observations of brood behavior after fledging. Females made 60% and 78% of total observed feedings of nestlings and fledglings, respectively. Parental provisioning rates increased with nestling age, and per‐nestling provisioning rates increased with brood size. Parental provisioning rates declined just before fledging, then increased just after fledging. Fledging times of individuals in broods were asynchronous and concentrated during the late afternoon and early evening. Males stopped caring for fledglings before females even though this species is single‐brooded, with some late‐season broods being abandoned by males. Broods spent the first three weeks after fledging within 400 m of nests, after which they began to disperse. Most aspects of the breeding biology of Cordilleran Flycatchers in our study, including the duration of nestling and fledging periods, female‐dominated provisioning, and movement patterns of fledglings, were similar to those of other Empidonax species. However, the times when young fledged were not concentrated in the morning as reported in most other songbirds, and this result warrants additional study of the timing of fledging in ecologically and taxonomically similar species. The increased per‐nestling provisioning rate with increasing brood size was unexpected, and additional study is needed to determine if this increase results from a trade‐off between adult annual survival and productivity favoring increased provisioning of young in larger broods, or from the existence of high‐quality individuals where larger clutches and higher provisioning rates are linked.     

Foraging behavior of Cerulean Warblers during the breeding and non‐breeding seasons: evidence for the breeding currency hypothesis

Birds require additional resources for raising young, and the breeding currency hypothesis predicts that insectivorous species exploit large soft‐bodied prey during the breeding season, but shift to small, likely hard‐bodied, prey during the non‐breeding season. To test this hypothesis, we examined prey use by Cerulean Warblers (Setophaga cerulea), foliage‐gleaning Nearctic‐Neotropical migrants, during the breeding and non‐breeding seasons. We collected data on foraging behavior during the breeding season (including observations of prey items fed to young) in upland mixed‐oak forest in southeastern Ohio in 2009 and 2010 and, during the non‐breeding season, in shade coffee in the Cordillera de Merida, Venezuela, in 2008–2009. Cerulean Warblers captured 7% more large prey (visible prey extending beyond the bill) during the breeding than the non‐breeding season, but foraged at similar rates during both seasons. Large, soft‐bodied prey appeared to be especially important for feeding young. We found that adults delivered large prey on >50% of provisioning visits to nests and 69% of identifiable large prey fed to nestlings were greenish larvae (likely Lepidoptera or caterpillars) that camouflage against leaves where they would tend to be captured by foliage‐gleaning birds. Availability of specific taxa appeared to influence tree species foraging preferences. As reported by other researchers, we found that Cerulean Warblers selected trees in the genus Carya for foraging and our examination of caterpillar counts from the central Appalachian Mountains (Butler and Strazanac 2000 ) showed that caterpillars with greenish coloration, especially Baileya larvae, may be almost twice as abundant on Carya than Quercus. Our results provide evidence for the breeding currency hypothesis, and highlight the importance of caterpillars to a foliage‐gleaning migrant warbler of conservation concern.     

Prey selection and foraging performance of breeding Great Tits Parus major in relation to food availability

We studied the nestling diet and the foraging performance of Great Tits in relation to prey abundance in the field. Numerous experimental studies present data on foraging decisions in captive Great Tits. Little is, however, known about prey selection in the field in relation to the food available and the consequences this has for the food delivery rate to nestlings. Since the foraging performance of the parents is one of the main determinants of fledging weight and juvenile survival, foraging behaviour is an important part of Great Tit reproduction. During the early breeding season up to 75% of the prey biomass delivered to the nestlings were spiders, which is in contrast with other studies. Only when caterpillars reached a size of 10–12 mg (approximately the average size of the spiders caught at that time) did the Great Tits change their preferences and 80–90% of the delivered prey masses were caterpillars, as reported by other authors. This 'switching' between prey occurred within a few days. It was not related to the changes in abundance but to size of caterpillars. The rate at which caterpillars were delivered to the nestlings (in mg/nestling/h) was strongly correlated with the caterpillar biomass available (in mg/m of branches) and nestling growth rate was significantly influenced by the mass of available caterpillars. The results provide evidence why perfect timing of breeding is so important for the Great Tit, and contribute to the understanding of the causal link between food supply, growth and breeding success.     

Feeding ecology and life history variation of the blue tit in Mediterranean deciduous and sclerophyllous habitats

Summary High variation in laying date and clutch size of the blue tit between a Mediterranean mixed habitat on the mainland, southern France, and a sclerophyllous habitat on the island of Corsica is hypothesized to be related to differences in the food supply. The diet of the nestlings and feeding frequencies were studied using camera nestboxes and electronic chronographs. Food items brought to the nestlings were much more diverse on Corsica than on the mainland, including many fewer caterpillars and a wider range of taxa. However, when expressed as a volume index, prey items were on average larger on Corsica than on the mainland. Feeding frequencies were significantly lower on Corsica. A good correlation was found in both habitats between laying date and the caterpillar peak date, although both the leafing development of oaks and the peak of abundance of caterpillars occurred 3 weeks later in the Corsican sclerophyllous trees than in the mainland deciduous ones. Differences in the feeding ecology of tits between the two habitats are discussed in the light of the evergreen habit, which means that only 30% of leaves are available for phyllophagous insects instead of 100% in deciduous trees. the combination of a late and low food supply in evergreen trees is the best explanation for the differences in breeding traits betwen the two populations.     

Nestling diet variation in an insular Mediterranean population of blue tits Parus caeruleus: effects of years,territories and individuals

Mediterranean evergreen forests of Corsica are characterized by relatively high species diversity of arthropods with low population densities. Food is never superabundant for Corsican blue tits Parus caeruleus. This study focused on the composition of the food of blue tit nestlings and especially on two main components, caterpillars and spiders. The nestling diet was studied for two years using 8-mm cameras that automatically took photographs of adult birds with food. The diet was composed of c. 50% caterpillars and c. 30% spiders. There were between-year and between-individual differences in these proportions. In both years of the study the proportion of caterpillars declined during the course of the breeding season. Individual and time effects on prey sizes were also observed. Pairs and individuals were fairly constant in the proportions of prey over the feeding period. Different food items were not brought in runs. These findings suggest that strong food limitation exists on Corsica, which can considerably influence life-history traits of the blue tit.     

Caterpillar abundance in the territory affects the breeding performance of great tit Parus major minor

The effects of caterpillar food supply on the breeding performance of a population of the Japanese great tit Parus major minor were investigated. Since more than 90% of the food items in our study site were caterpillars living on trees, we estimated the food availability using 20 frass traps per hectare. The sampling error of this method was about 10% on average, which was accurate enough to detect differences between territories. Food abundance at laying in each territory affected the timing of egg laying. However, food amount after hatching was correlated with clutch size. No relationship was found between fledgling quality and food availability, probably because the effects of local variation in food abundance could be canceled out by parental effort such as extending the foraging area. There was a significant negative correlation between the length of the nestling period and food availability. We suggest that parent tits decide the timing of fledging at the point where two factors, predation risk before fledging and additional improvement of nestling quality, are balanced. Food availability just after fledging affected the length of post-fledging parental care; it seems that fledglings in “poor” territories would have had difficulty in finding food and hence needed to depend on their parents longer than those in “rich” territories. Received: 10 June 1997 / Accepted: 29 December 1997     

Plumage yellowness predicts foraging ability in the blue tit Cyanistes caeruleus

Carotenoid‐based coloration in adult birds has been often regarded as an honest signal of individual quality. However, few studies have demonstrated a link between carotenoid display and the quantity or quality of resources provided to the offspring. The present study investigated the expression of a carotenoid‐based ornament, the breast plumage yellowness of the blue tit Cyanistes caeruleus, in relation to the level of parental provisioning effort and the amount of carotenoid‐rich prey provided to the young. The study was conducted in two forest types (evergreen and deciduous), which also allowed an exploration of the possible existence of habitat effects on the coloration of breeding birds. It was found that plumage colour intensity (carotenoid chroma) correlated positively with nestling provisioning rates of both males and females, supporting the good parent hypothesis. In addition, carotenoid chroma was positively related with the proportion of Lepidoptera larvae brought to the nest in both sexes. Female but not male coloration was positively linked to breeding success (proportion of fledged young). Nestling coloration did not correlate with that of their parents, nor the frequency with which they were fed. Hue and lightness of nestling's plumage correlated positively with body mass and tarsus length, respectively. The results obtained in the present study indicate that ventral plumage coloration in blue tits may advertise the ingested carotenoids (carotenoid foraging ability) and also their overall parental quality in terms of nestling provisioning rates. This suggests that plumage yellowness can be used as an indicator of foraging ability in this species. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 418–429.     

Seasonal variation in diet quality: antioxidants,invertebrates and blue tits Cyanistes caeruleus

Breeding success is often dictated by the degree to which parents can synchronize the maximum food requirements of offspring to the peak in abundance of invertebrate prey. Less studied is how the nutritional quality of individual diet items impacts on breeding. In the present study, we assessed the abundance and antioxidant concentrations of arboreal arthropods from oak woodland and provisioning behaviour of the blue tit Cyanistes caeruleus. Dietary antioxidants are important during development because they defend against oxidative stress. Operophtera caterpillars, Erannis caterpillars, and spiders contained significantly different levels of individual carotenoids and α‐tocopherol. Concentrations of lutein and β‐carotene in Operophtera caterpillars did not vary seasonally, although concentrations of zeaxanthin declined and α‐tocopherol increased with date. Blue tit broods hatched later in the season received significantly fewer caterpillars and more spiders per chick compared to earlier broods. Reflecting changes in prey composition, blue tit nestling plasma showed decreases in zeaxanthin and increases in α‐tocopherol with date. Thus, processes that shift the timing of breeding in birds and/or prey composition are likely to alter antioxidant intake and thus potentially influence the oxidative stress status of animals. The data obtained in the present study suggest a mechanism by which environmental change as a result of human activities could influence the health and fitness of individuals in natural populations. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 708–717.     

Sea surface temperatures and behavioural buffering capacity in thin-billed prions Pachyptila belcheri: breeding success, provisioning and chick begging

Seabirds are important predators in marine ecosystems and are commonly used to monitor the productivity of their marine environments. However, different measures of seabird breeding success differ in their sensitivity to environmental conditions. Here, we present an analysis of provisioning rates and chick growth as well as hatching and fledging success, in thin-billed prions Pachyptila belcheri at New Island, Falkland Islands between 2003 and 2005 and relate these patterns to ocean climate. During the study period, SST were rising within and between breeding seasons and were negatively correlated with provisioning frequencies of thin-billed prions. Chick mass was reduced and begging intensities increased at low feeding frequencies, but overall breeding success and fledging success were not affected, because most chicks in survived to fledging despite poor provisioning rates. Chick numbers and survival therefore may not be sensitive indicators of environmental conditions in all seabird species and ranges of food abundance. Monitoring behavioural buffering mechanisms, such as feeding rates, may be more effective in some ranges of food abundance and time scales and provide an earlier warning of ecological change. As a novel technique, the use of begging intensities as indicator of chick body condition is proposed in species where repeated handling causes disturbance. The present data suggest that begging rates can serve as a non-invasive method to monitor chick condition.     

Diet flexibility in a declining long‐distance migrant may allow it to escape the consequences of phenological mismatch with its caterpillar food supply

Climate‐driven shifts in prey phenology may lead to asynchrony with the timing of peak resource requirements of their predators, leading to a reduction in productivity and population declines. Migrant species that cannot adjust their arrival times may be particularly at risk, especially those that breed in seasonal environments and for which a temporarily super‐abundant prey source is important, such as insectivorous passerine birds that take advantage of the seasonal flush of caterpillars to feed their young. We assess whether population declines of the trans‐Saharan migratory Wood Warbler Phylloscopus sibilatrix are likely to have been caused by phenological mismatch. We measured seasonal invertebrate biomass and various fitness parameters, including the timing of breeding and breeding success, in two time periods: 1982–1984, prior to the species’ decline in the UK, and 2009–2011, as the reduction in numbers continued. Although birds bred on average a week earlier in 2009–2011 than in 1982–1984, this was not adequate to track the more rapid advancement of peak caterpillar biomass, which advanced by 12 days and was closely correlated with spring temperatures. Moreover, although caterpillars were the dominant prey fed to nestlings, there was only limited evidence that productivity was positively related to caterpillar biomass in the environment. Considering only successful nests, synchrony with the food peak did not produce heavier nestlings and had only a small positive effect on fledging success, although there was a seasonal decline in productivity when all nests were considered. We conclude that the lack of a marked effect of the observed mismatch is due to Wood Warblers’ generalist diet, enabling them to breed successfully on prey other than caterpillars. Although other studies have demonstrated that climate‐driven asynchrony of predator and prey populations can have impacts on avian demography, this study highlights the importance of investigating the generality of those findings.     

Caterpillar biomass depends on temperature and precipitation,but does not affect bird reproduction

Complex changes in phenological events appear as temperatures are increasing: In deciduous forests bud burst, hatching of herbivorous caterpillars, egg laying and nestling time of birds when feeding chicks on caterpillars, may differentially shift into early season and alter synchronization. If timing of bird reproduction has to match with short periods of food availability, phenological mismatch could negatively affect reproductive success. Using a unique empirical approach along an altitudinal temperature gradient, we firstly asked whether besides temperature, also precipitation and leaf phenology interplay and affect caterpillar biomass, since impacts of rainfall on caterpillars have been largely neglected so far. Secondly, we asked whether abundance of caterpillars and thereby body mass of great tit nestlings, which are mainly fed with caterpillars, vary along the altitudinal temperature gradient. We demonstrated that next to temperature also precipitation and leaf phenology affected caterpillar biomass. In our beech forest, even along altitudes, caterpillars were available throughout the great tit breeding season but in highly variable amounts. Our findings revealed that although timing of leaf phenology and great tit breeding season were delayed with decreasing temperature, caterpillars occurred synchronously and were not delayed according to altitude. However, altitude negatively affected caterpillar biomass, but body mass of fledglings at high altitude sites was not affected by lower amounts of caterpillar biomass. This might be partially outweighed by larger territory sizes in great tits.     

Is the blue tit falling into an ecological trap in Argentine ant invaded forests?

Because insectivorous birds must evaluate resources for reproduction before settling into a breeding habitat, they can fall into an ecological trap if informative cues about habitat suitability become dissociated from their actual yield. Given their potential to affect ecological networks, invasive ant species are potential candidates for triggering such ecological traps. We combined observational and experimental approaches to examine whether the variation in food supply for nestlings resulting from the invasion of the Argentine ant, Linepithema humile, had any influence on the breeding ecology of the blue tit, Cyanistes caeruleus, an insectivorous foliage-gleaner. We investigated the effects of the ant invasion on breeding performance (nesting success, clutch size, brood size and breeding success) and offspring quality (body size and condition, developmental stability and plumage colour) in replicated Mediterranean forest areas over a period of 3 years. There was no evidence that the reduction in caterpillar availability resulting from the invasion had a concurrent negative effect on the blue tit’s ability to successfully rear nestlings in optimal conditions, at least as measured here. Although the raw figures suggest an increased level of nutritional stress in blue tits breeding in invaded forests, the data analyses showed no significant alterations in terms of productivity or offspring fitness. The reproductive performance of the blue tit has been shown to be remarkably resilient to the Argentine ant-mediated food shortage, either because the prey reduction following the invasion did not reach a critical threshold or because of compensatory activity by the progenitors. We cannot conclusively reject an ecological trap triggered by the ant invasion on blue tits, since neither fledgling recruitment nor the prospective survival of parents were assessed. Even though we could not confirm short-term consequences of the Argentine ant invasion on blue tit reproductive fitness, the long-term bottom-up effects of the invasion remain unknown and should not be ruled out.     

Parental food provisioning is related to nestling stress response in wild great tit nestlings: implications for the development of personality

Background

Variation in early nutrition is known to play an important role in shaping the behavioural development of individuals. Parental prey selection may have long-lasting behavioural influences. In birds foraging on arthropods, for instance, the specific prey types, e.g. spiders and caterpillars, matter as they have different levels of taurine which may have an effect on personality development. Here we investigated how naturally occurring variation in the amounts of spiders and caterpillars, provisioned to nestlings at day 4 and 8 after hatching, is related to the response to handling stress in a wild passerine, the great tit (Parus major). Broods were cross-fostered in a split-brood design allowing us to separate maternal and genetic effects from early rearing effects. Adult provisioning behaviour was monitored on day four and day eight after hatching using video recordings. Individual nestlings were subjected to a handling stress test at an age of 14 days, which is a validated proxy for exploratory behaviour as an adult.

Results

Variation in handling stress was mainly determined by the rearing environment. We show that, contrary to our predictions, not the amount of spider biomass, but the amount of caterpillar biomass delivered per nestling significantly affected individual performance in the stress test. Chicks provisioned with lower amounts of caterpillars exhibited a stronger stress response, reflecting faster exploratory behaviour later on in life, than individuals who received larger amounts of caterpillars.

Conclusions

These results suggest that natural variation in parental behaviour in wild birds modulates the developmental trajectories of their offspring's personality via food provisioning. Since parental provisioning behaviour might also reflect the local environmental conditions, provisioning behaviour may influence how nestlings respond to these local environmental conditions.