Great Tit Parus Major Population Research Articles

Breeding parameters of a Great Tit (Parus major) population on the island of Cyprus compared to other Mediterranean populations

Many studies on the Great Tit, Parus major, have been conducted within the Mediterranean Basin, especially on its reproductive biology, forest habitat interactions, and nutritional resources. However, to date no detailed scientific research on these issues has been conducted on the island of Cyprus with a general paucity in research on this topic from the Eastern Mediterranean. Island species, through reduced gene flow and general isolation often evolve reproductive differences from their mainland relatives. As Cyprus is the third largest island in the Mediterranean and the most easterly positioned it provides an ideal opportunity compare the breeding biology of this island population with those from the wider Mediterranean region. This study aimed to gather some baseline reproductive data for Great Tits breeding in Cyprus. This data was then used to compare the reproductive outputs of Cypriot Great Tits from the sub species Parus major aphrodite with other Mediterranean island populations. This study was performed over a 3-year period during the breeding seasons of 2016-2018 in four evergreen forest areas. Mean clutch size was 7.7 eggs and hatching success 76.0%. The mean weight of the young was 16.3 g and mean tarsus length 18.7 mm. Significant differences were found between the years of the study, both in mean clutch size (p<0.0001) and the day of the first egg (p<0.0001), with 2016 producing the largest clutches and clutches being laid earlier than the subsequent two years. The start of the breeding season of the Great Tit was considerably earlier in Cyprus compared to other Mediterranean regions.

Metabolic adjustments to winter severity in two geographically separated great tit (Parus major) populations.

Understanding the potential limits placed on organisms by their ecophysiology is crucial for predicting their responses to varying environmental conditions. A main hypothesis for explaining avian thermoregulatory mechanisms is the aerobic capacity model, which posits a positive correlation between basal (basal metabolic rate [BMR]) and summit (Msum) metabolism. Most evidence for this hypothesis, however, comes from interspecific comparisons, and the ecophysiological underpinnings of avian thermoregulatory capacities hence remain controversial. Indeed, studies have traditionally relied on between-species comparisons, although, recently, there has been a growing recognition of the importance of intraspecific variation in ecophysiological responses. Therefore, here, we focused on great tits (Parus major), measuring BMR and Msum during winter in two populations from two different climates: maritime-temperate (Gontrode, Belgium) and continental (Zvenigorod, Russia). We tested for the presence of intraspecific geographical variation in metabolic rates and assessed the predictions following the aerobic capacity model. We found that birds from the maritime-temperate climate (Gontrode) showed higher BMR, whereas conversely, great tits from Zvenigorod showed higher levels of Msum. Within each population, our data did not fully support the aerobic capacity model's predictions. We argued that the decoupling of BMR and Msum observed may be caused by different selective forces acting on these metabolic rates, with birds from the continental-climate Zvenigorod population facing the need to conserve energy for surviving long winter nights (by keeping their BMR at low levels) while simultaneously being able to generate more heat (i.e., a high Msum) to withstand cold spells.

The genomics of adaptation to climate in European great tit (Parus major) populations.

The recognition that climate change is occurring at an unprecedented rate means that there is increased urgency in understanding how organisms can adapt to a changing environment. Wild great tit (Parus major) populations represent an attractive ecological model system to understand the genomics of climate adaptation. They are widely distributed across Eurasia and they have been documented to respond to climate change. We performed a Bayesian genome-environment analysis, by combining local climate data with single nucleotide polymorphisms genotype data from 20 European populations (broadly spanning the species' continental range). We found 36 genes putatively linked to adaptation to climate. Following an enrichment analysis of biological process Gene Ontology (GO) terms, we identified over-represented terms and pathways among the candidate genes. Because many different genes and GO terms are associated with climate variables, it seems likely that climate adaptation is polygenic and genetically complex. Our findings also suggest that geographical climate adaptation has been occurring since great tits left their Southern European refugia at the end of the last ice age. Finally, we show that substantial climate-associated genetic variation remains, which will be essential for adaptation to future changes.

Variation in DNA Methylation in Avian Nestlings Is Largely Determined by Genetic Effects.

As environmental fluctuations are becoming more common, organisms need to rapidly adapt to anthropogenic, climatic, and ecological changes. Epigenetic modifications and DNA methylation in particular provide organisms with a mechanism to shape their phenotypic responses during development. Studies suggest that environmentally induced DNA methylation might allow for adaptive phenotypic plasticity that could last throughout an organism's lifetime. Despite a number of studies demonstrating environmentally induced DNA methylation changes, we know relatively little about what proportion of the epigenome is affected by environmental factors, rather than being a consequence of genetic variation. In the current study, we use a partial cross-foster design in a natural great tit (Parus major) population to disentangle the effects of common origin from common rearing environment on DNA methylation. We found that variance in DNA methylation in 8,315 CpG sites was explained by a common origin and only in 101 by a common rearing environment. Subsequently, we mapped quantitative trait loci for the brood of origin CpG sites and detected 754 cis and 4,202 trans methylation quantitative trait loci, involving 24% of the CpG sites. Our results indicate that the scope for environmentally induced methylation marks independent of the genotype is limited and that the majority of variation in DNA methylation early in life is determined by genetic factors instead. These findings suggest that there may be little opportunity for selection to act on variation in DNA methylation. This implies that most DNA methylation variation likely does not evolve independently of genomic changes.

No Apparent Immediate Reproductive Costs of Overlapping Breeding and Moult in a Mediterranean Great Tit Population.

Some phenological events in birds, such as breeding and moulting, are being affected by rising temperatures due to global warming, and many species have undergone temporary changes in these energetically demanding phases that are often separated in time. This has led to an increased overlap between breeding and moulting in some populations. This overlap causes conflicts in resource allocation and may impose fitness costs that could affect immediate reproductive performance. We tested whether this occurs in a great tit (Parus major) population in eastern Spain. In 71% of 390 pairs, in which both parents were captured during the period of overlap between moulting and breeding, at least one parent was moulting when feeding the chicks of its second brood. Later breeders were more likely to overlap breeding and moulting, and when both parents overlapped, clutch size was smaller, fewer eggs hatched and fewer fledglings in poorer body condition were produced. Some results were intermediate when only one parent moulted. However, all these differences between moulting and non-moulting pairs disappeared when the seasonal trend in reproductive parameters was taken into account, as moulting birds bred later and reproductive performance decreased seasonally. Therefore, the overlap of breeding and moulting does not impose additional reproductive costs in this population.

Differences in feather structure between urban and forest great tits: constraint or adaptation?

Urbanization is one of the strongest habitat transforming processes today that has resulted in changes in the ecological conditions for wild populations. In birds, the limitation of natural food sources and a warmer microclimate in cities can potentially influence the development and functioning of the plumage that may have important fitness consequences. Despite its potential significance, the plumage structure of urban birds are largely unexplored and it is unclear whether and how they respond to urban ecological processes such as different constraints and selection pressures. In this study, we compared several structural properties of contour, primary and tail feathers between two forest and two urban great tit Parus major populations. Our results show that the urban environment affects only a few structural properties of feathers and only in the plumage of first‐year birds. We found that both the plumulaceous and the pennaceous parts of their contour feathers are longer and have lower barb density in the urban than in the forest habitat. We also found that the primaries of first‐year birds have narrower rachis and higher barbule density in the cities than in the forests, but there were no differences in other wing feather traits and in any tail feather traits between habitats. We did not find differences in the feather structure of urban versus forest adult birds. The habitat differences in first‐year birds may indicate nutritional constraints or the effects of the warmer microclimate of the urban environment. These differences seem to disappear completely in adulthood that can be explained by the selective mortality of first‐year birds, or by adults being less sensitive than first‐year birds to environmental effects during their moult.

Effect of Human Disturbance on Bird Telomere Length: An Experimental Approach

Human recreational activities increase worldwide in space and frequency leading to higher rates of encounter between humans and wild animals. Because wildlife often perceive humans as predators, this increase in human disturbance may have negative consequences for the individuals and also for the viability of populations. Up to now, experiments on the effects of human disturbance on wildlife have mainly focused on individual behavioral and stress-physiological reactions, on breeding success, and on survival. However, the effects on other physiological parameters and trans-generational effects remain poorly understood. We used a low-intensity experimental disturbance in the field to explore the impacts of human disturbance on telomere length in great tit (Parus major) populations and found a clear effect of disturbance on telomere length. Adult males, but not females, in disturbed plots showed shorter telomere lengths when compared to control plot. Moreover, variation in telomere length of adult great tits was reflected in the next generation, as we found a positive correlation between telomere length of the chicks and of their fathers. Given that telomere length has been linked to animal lifespan, our study highlights that activities considered to be of little concern (i.e., low levels of disturbance) can have a long-lasting impact on the physiology and survival of wild animals and their next generation.

Double-brooding and annual breeding success of great tits in urban and forest habitats.

Urban areas differ from natural habitats in several environmental features that influence the characteristics of animals living there. For example, birds often start breeding seasonally earlier and fledge fewer offspring per brood in cities than in natural habitats. However, longer breeding seasons in cities may increase the frequency of double-brooding in urban compared with nonurban populations, thus potentially increasing urban birds' annual reproductive output and resulting in lower habitat difference in reproductive success than estimated by studies focusing on first clutches only. In this study, we investigated 2 urban and 2 forests great tit Parus major populations from 2013 to 2019. We compared the probability of double-brooding and the total number of annually fledged chicks per female between urban and forest habitats, while controlling for the effects of potentially confounding variables. There was a trend for a higher probability of double-brooding in urban (44% of females) than in forest populations (36%), although this was not consistent between the 2 urban sites. Females produced significantly fewer fledglings annually in the cities than in the forest sites, and this difference was present both within single- and double-brooded females. Furthermore, double-brooded urban females produced a similar number of fledglings per season as single-brooded forest females. These results indicate that double-brooding increases the reproductive success of female great tits in both habitats, but urban females cannot effectively compensate in this way for their lower reproductive output per brood. However, other mechanisms like increased post-fledging survival can mitigate habitat differences in reproductive success.

A higher incidence of moult–breeding overlap in great tits across time is linked to an increased frequency of second clutches: a possible effect of global warming?

The rise of temperatures due to global warming is related to a lengthening of the breeding season in many bird species. This allows more pairs to attempt two clutches within the breeding season, thus finishing their breeding activity later in the season and therefore potentially overlapping these with post–breeding moult. We tested whether this occurred in two Spanish great tit Parus major populations. The proportion of pairs laying second clutches increased from 1 % to 32 % over the study period in one of the populations (Sagunto, 1995–2019), while it did not change in the other (Quintos, 2006–2019; mean 5 %). We did not find any temporal trend for moult start date of late–breeding birds in any population. The proportion of individuals of both sexes that overlapped moult and breeding increased in Sagunto. For this latter population, sex and age, but not clutch type, contributed to the variability in the probability of overlapping in late–breeding individuals, this being higher for first–year males and lower for older females.

Deconstructing incubation behaviour in response to ambient temperature over different timescales

Avian embryos need a stable thermal environment to develop optimally, while incubating females need to allocate time to self‐maintenance off the nest. In species with female‐only incubation, eggs are exposed to ambient temperatures that usually cool them down during female absences. The lower the ambient temperature the sooner females should return to re‐warm the eggs. When incubation constraints ease at increasing ambient temperatures, females respond by increasing either incubation effort or self‐maintenance time. These responses are population‐dependent even within the same species; but it is uncertain whether they are caused by local environmental conditions or they are an artefact from limited datasets, different methodological approaches or the timescale over which incubation behaviour is measured. In this study, we collected incubation data from three Mediterranean great tit Parus major populations during three consecutive years. We measured the duration of each off‐ and on‐bout event, used these variables to compute nest attentiveness at three different timescales (full incubation, daily and hourly periods) and assessed the impact of ambient temperature on bout duration and nest attentiveness. We found that females maximized on‐bout duration at different local temperatures, ranging from 10 to 20°C; but lengthened off‐bouts linearly across a range of 0–38°C in all three populations. These local differences translated into opposite linear nest attentiveness patterns at the full incubation scale: Females increased either incubation effort, longest on‐bouts between 15 and 20°C or self‐maintenance time, longest on‐bouts at 10°C. It was at daily and hourly periods when we detected non‐linear nest attentiveness patterns, as expected from on‐bout duration, peaking at different local ambient temperatures. Females first increased incubation effort up to a certain temperature value and then increased self‐maintenance time at the highest ambient temperatures. Further research is needed to understand which factors are behind the turning points from one behaviour to the other.

Impacts of ambient temperature and clutch size on incubation behaviour onset in a female‐only incubator songbird

Ambient temperature is assumed to be the major cue used by passerines to synchronize their laying and hatching dates to the expected peak of prey availability. While laying eggs, females are still able to fine‐tune their hatching date following increasing or decreasing patterns of ambient temperature, mostly via changes in incubation onset. The onset of incubation behaviour in relation to the laying sequence could have later consequences for the duration of the incubation period and the extent of hatching asynchrony. Clutch size is also known to affect incubation patterns and might therefore condition potential responses to changing temperatures. In this study we assessed the association of ambient temperature and clutch size with the onset of four different incubation behaviours: partial and full nocturnal incubation, and partial and full diurnal incubation. We also evaluated how the onset and duration of each incubation behaviour might predict the duration of diurnal full incubation and the extent of hatching asynchrony. To achieve our aims, we monitored incubation behaviour using temperature data loggers during the egg‐laying period in three Mediterranean Great Tit Parus major populations in three consecutive years. Our results showed that increasing temperatures were related to an advance of diurnal partial incubation, but not its duration or the onset of full incubation behaviour. We did not find any effect of ambient temperatures on nocturnal incubation. However, females lengthened nocturnal partial incubation and delayed the onset of nocturnal full and diurnal partial incubation when laying larger clutches. Longer diurnal incubation before clutch completion was associated with greater hatching asynchrony. Moreover, longer diurnal partial incubation shortened the duration of the full incubation period. In conclusion, increasing ambient temperatures during the egg‐laying period advanced diurnal partial incubation, indirectly shortening the full incubation period and increasing hatching asynchrony.

Providing urban birds nutritious food to feed chicks reduces urban versus rural breeding success disparities.

In Focus: Seress, G, Sándor, K, Evans, KL, Liker, A. (2020) Food availability limits avian reproduction in the city: An experimental study on great tits Parus major . Journal of Animal Ecology. 89. Animals in urban areas face a multitude of stressors, but how each stressor impacts survival and fitness can be difficult to disentangle. We need experimental manipulations of suspected stressors to examine causal relationships with traits of free-living urban and rural animals. In this issue, Seress and colleagues take on an intensive experimental approach to test whether one potential stressor-limited natural food sources in cities-can explain reduced avian reproductive success in urban areas. They employ a full factorial design, including both food supplemented and control broods in both urban and forest great tit Parus major populations. The findings are clear. Reduced food availability is the key factor limiting urban offspring growth and survival, at least in this well-studied species. Indeed, the extra insects fed to urban chicks greatly reduced the significant differences in survival rates and body sizes between urban and forest broods. The findings are also sobering. Urban insect populations would need to more than double to erase differences in reproductive success between urban and rural bird populations, an unlikely outcome with the ever-increasing urbanization of habitats.

It takes two: Heritable male effects on reproductive timing but not clutch size in a wild bird population.

Within-population variation in the traits underpinning reproductive output has long been of central interest to biologists. Since they are strongly linked to lifetime reproductive success, these traits are expected to be subject to strong selection and, if heritable, to evolve. Despite the formation of durable pair bonds in many animal taxa, reproductive traits are often regarded as female-specific, and estimates of quantitative genetic variation seldom consider a potential role for heritable male effects. Yet reliable estimates of such social genetic effects are important since they influence the amount of heritable variation available to selection. Based on a 52-year study of a nestbox-breeding great tit (Parus major) population, we apply "extended" bivariate animal models in which the heritable effects of both sexes are modeled to assess the extent to which males contribute to heritable variation in seasonal reproductive timing (egg laying date) and clutch size, while accommodating the covariance between the two traits. Our analyses show that reproductive timing is a jointly expressed trait in this species, with (positively covarying) heritable variation for laydate being expressed in both members of a breeding pair, such that the total heritable variance is 50% larger than estimated by traditional models. This result was robust to explicit consideration of a potential male-biased environmental confound arising through sexually dimorphic dispersal. In contrast to laydate, males' contribution to heritable variation in clutch size was limited. Our study thus highlights the contrasting extent of social determination for two major components of annual reproductive success, and emphasizes the need to consider the social context of what are often considered individual-level traits.

Longitudinally monitored lifetime changes in blood heavy metal concentrations and their health effects in urban birds

Urban heavy metal pollution can impair the health of humans and other organisms inhabiting cities. While birds are suggested as one of the appropriate bioindicators for essential and non-essential trace element monitoring, the process of particular elements' accumulation in blood and its possible adverse health effects during ageing of individuals remain unexplored. We have investigated lifetime changes in blood lead (Pb), cadmium (Cd), arsenic (As) and zinc (Zn) concentrations and searched for links to health-related traits in sub-urban free-living great tit (Parus major) population monitored over a long period of time. The blood As concentrations were under the limit of detection in most samples. The blood Pb levels showed a non-linear relationship to individuals age, where the highest Pb concentrations were measured in nestlings and in a very small group of highly senescent birds (over 7 years old), while no clear trend was observed for the majority of the adult age stages. No age-related patterns were found for blood Cd or Zn concentrations. The positive relationship between date of capture and blood Cd and Zn levels may reflect seasonal changes in diet composition. We did not reveal any anaemia-like conditions (decreased total erythrocyte count or increased immature erythrocyte count) in relation to blood heavy metal concentrations in the investigated birds. Total leukocyte counts, heterophil/lymphocyte (H/L) ratio and total heterophil and lymphocyte counts increased with increasing Pb, Cd and Zn concentrations in blood. This study demonstrates the suitability of avian blood for actual heavy metal spatial and temporal biomonitoring even in situations when the precise age of the individuals remains unknown.

Extreme temperature drop alters hatching delay, reproductive success, and physiological condition in great tits

It has been suggested that extreme weather events may be treated as natural experiments that may unravel the mechanisms by which birds adjust their phenology and breeding parameters to environmental variability. In 2017, a sudden and heavy drop of temperatures for several days affected many European bird populations. This event occurred during the laying–early incubation period in the great tit (Parus major) population in central Poland, causing a large delay in hatching and had sustained reproductive consequences. This cold snap occurring once breeding activity had already started in 2017 was followed by the warm and invariable breeding season of 2018. This natural experiment had an essential influence on great tit reproductive parameters. We found a significant difference in hatching date, number of fledglings, hatching success, and fledging success between 2017 and 2018. In 2017, there were about two fledglings per nest fewer than in 2018. Fledging success was positively associated with hatching delay in 2017, while the relation was negative in 2018. Hatching success differed significantly between both years, being higher in 2018. Mean level of hemoglobin (used as index of body, physiological condition) in the blood of nestling great tits was higher in 2017 than in 2018. We argue that the moment of hatching may be (at least in some years) more tightly related to the moment of maximum food demand of tit nestlings than the traditionally used laying date. Also in extreme years, phenotypic plasticity of hatching delays may be insufficient to adjust the timing of breeding to the upcoming extreme weather events. Further examining its limits may be an important goal for future research.

Effect of age, haemosporidian infection and body condition on pair composition and reproductive success in Great Tits Parus major

Several factors shape lifetime reproductive success, including genetic background, body condition, environmental conditions and ecological interactions such as parasitism. Adults often show higher reproductive success than their young conspecifics, especially in long‐lived bird species, and this may be explained by the cumulative effects of an increase in reproductive experience and the selection of high‐quality individuals from one year to the next. To test whether this pattern also exists in short‐lived bird species, we used 13 years of monitoring data from two Great Tit Parus major populations. The effects of male and female age on several reproductive parameters were analysed in 419 pairs of Great Tits, while accounting for body condition and infection by haemosporidian parasites. Reproductive success was mainly affected by the age‐class of males. Pairs containing a sub‐adult male fledged one‐third fewer chicks than pairs containing an adult male. The difference was not caused by variation in male fertility but could have been caused by better parental care provided by adult birds. In addition to lower reproductive success, first‐year males also had reduced access to mating compared with adult males, suggesting an avoidance of sub‐adult males by females. Nestling body condition was positively correlated with parental body condition, and the body condition of male and female members of breeding pairs was positively correlated. Finally, the number of fledged chicks was mainly affected by the infection status of males. These results temper our previously published results showing an effect of infection on Great Tit reproduction regardless of their sex. In our previous study, and as in most cases, the status of the partner was not taken into account and we show here that this is essential because it can lead to a biased interpretation of the results.

A high-density SNP chip for genotyping great tit (Parus major) populations and its application to studying the genetic architecture of exploration behaviour.

High-density SNP microarrays ("SNP chips") are a rapid, accurate and efficient method for genotyping several hundred thousand polymorphisms in large numbers of individuals. While SNP chips are routinely used in human genetics and in animal and plant breeding, they are less widely used in evolutionary and ecological research. In this article, we describe the development and application of a high-density Affymetrix Axiom chip with around 500,000 SNPs, designed to perform genomics studies of great tit (Parus major) populations. We demonstrate that the per-SNP genotype error rate is well below 1% and that the chip can also be used to identify structural or copy number variation. The chip is used to explore the genetic architecture of exploration behaviour (EB), a personality trait that has been widely studied in great tits and other species. No SNPs reached genomewide significance, including at DRD4, a candidate gene. However, EB is heritable and appears to have a polygenic architecture. Researchers developing similar SNP chips may note: (i) SNPs previously typed on alternative platforms are more likely to be converted to working assays; (ii) detecting SNPs by more than one pipeline, and in independent data sets, ensures a high proportion of working assays; (iii) allele frequency ascertainment bias is minimized by performing SNP discovery in individuals from multiple populations; and (iv) samples with the lowest call rates tend to also have the greatest genotyping error rates.

Haemosporidian infections affect antioxidant defences in great tits Parus major but are not related to exposure to aerial pollutants

SUMMARY Haemosporidian parasites are widespread in avian species and modulate their ecology, behaviour, life-history and fitness. The prevalence of these vector-transmitted parasites varies with host intrinsic and extrinsic factors, such as host resistance and behaviour, and habitat-related characteristics. In this study, we evaluated the prevalence of avian haemosporidians in great tit Parus major populations inhabiting two areas with different degrees of exposure to aerial emissions from pulp mills, to assess if this type of pollution impacted parasite prevalence. We also compared the physiological condition of infected and uninfected individuals. Haemosporidian infection prevalence was high (58%), varied seasonally, but was not associated with air pollution exposure. Fledged birds presented higher infection rates than nestlings and infected fledged birds had higher levels of blood glutathione peroxidase activity. These results allow us to infer that infection by blood parasites may activate antioxidant defences, possibly to protect the organism from the negative oxidative stress side-effects of immune activation against parasites. Because oxidative stress is one of the mechanisms responsible for ageing and senescence and may affect fitness, the relationship between parasitism and oxidative stress markers should be further investigated through studies that include experimental manipulation of infection in model organisms.

Density dependence in an age-structured population of great tits: identifying the critical age classes.

Classical approaches for the analyses of density dependence assume that all the individuals in a population equally respond and equally contribute to density dependence. However, in age-structured populations, individuals of different ages may differ in their responses to changes in population size and how they contribute to density dependence affecting the growth rate of the whole population. Here we apply the concept of critical age classes, i.e., a specific scalar function that describes how one or a combination of several age classes affect the demographic rates negatively, in order to examine how total density dependence acting on the population growth rate depends on the age-specific population sizes. In a 38-yr dataset of an age-structured great tit (Parus major) population, we find that the age classes, including the youngest breeding females, were the critical age classes for density regulation. These age classes correspond to new breeders that attempt to take a territory and that have the strongest competitive effect on other breeding females. They strongly affected population growth rate and reduced recruitment and survival rates of all breeding females. We also show that depending on their age class, females may differently respond to varying density. In particular, the negative effect of the number of breeding females was stronger on recruitment rate of the youngest breeding females. These findings question the classical assumptions that all the individuals of a population can be treated as having an equal contribution to density regulation and that the effect of the number of individuals is age independent. Our results improve our understanding of density regulation in natural populations.

Genetic variation in variability: Phenotypic variability of fledging weight and its evolution in a songbird population.

Variation in traits is essential for natural selection to operate and genetic and environmental effects can contribute to this phenotypic variation. From domesticated populations, we know that families can differ in their level of within-family variance, which leads to the intriguing situation that within-family variance can be heritable. For offspring traits, such as birth weight, this implies that within-family variance in traits can vary among families and can thus be shaped by natural selection. Empirical evidence for this in wild populations is however lacking. We investigated whether within-family variance in fledging weight is heritable in a wild great tit (Parus major) population and whether these differences are associated with fitness. We found significant evidence for genetic variance in within-family variance. The genetic coefficient of variation (GCV) was 0.18 and 0.25, when considering fledging weight a parental or offspring trait, respectively. We found a significant quadratic relationship between within-family variance and fitness: families with low or high within-family variance had lower fitness than families with intermediate within-family variance. Our results show that within-family variance can respond to selection and provides evidence for stabilizing selection on within-family variance.