Population Of Great Tits Research Articles

The Genetic Response of Forest Birds to Urbanization: Variability in the Populations of Great and Blue Tits

The Genetic Response of Forest Birds to Urbanization: Variability in the Populations of Great and Blue Tits

Extra-pair paternity and sperm length variation in a far northern Great Tit (Parus major) population

Extra-pair paternity and sperm length variation in a far northern Great Tit (Parus major) population

The great tit HapMap project: A continental-scale analysis of genomic variation in a songbird.

A major aim of evolutionary biology is to understand why patterns of genomic diversity vary within taxa and space. Large-scale genomic studies of widespread species are useful for studying how environment and demography shape patterns of genomic divergence. Here, we describe one of the most geographically comprehensive surveys of genomic variation in a wild vertebrate to date; the great tit (Parus major) HapMap project. We screened ca 500,000 SNP markers across 647 individuals from 29 populations, spanning ~30 degrees of latitude and 40 degrees of longitude - almost the entire geographical range of the European subspecies. Genome-wide variation was consistent with a recent colonisation across Europe from a South-East European refugium, with bottlenecks and reduced genetic diversity in island populations. Differentiation across the genome was highly heterogeneous, with clear 'islands of differentiation', even among populations with very low levels of genome-wide differentiation. Low local recombination rates were a strong predictor of high local genomic differentiation (FST), especially in island and peripheral mainland populations, suggesting that the interplay between genetic drift and recombination causes highly heterogeneous differentiation landscapes. We also detected genomic outlier regions that were confined to one or more peripheral great tit populations, probably as a result of recent directional selection at the species' range edges. Haplotype-based measures of selection were related to recombination rate, albeit less strongly, and highlighted population-specific sweeps that likely resulted from positive selection. Our study highlights how comprehensive screens of genomic variation in wild organisms can provide unique insights into spatio-temporal evolutionary dynamics.

Effects of an extreme weather event over reproduction and survival of Great Tits (Parus major) in eastern Spain

The frequency of extreme climatic and weather events has increased since 1950 due to global changes induced by human influence. These events can have significant impacts at the individual, population, and community levels across various taxonomic groups. They can be particularly detrimental to bird populations during their breeding season, affecting specific breeding parameters. This study originated from an exceptionally rare breeding season of a resident Great Tit population recorded in spring 2002 when an unusual mortality of nestlings was observed. We quantified weather conditions (temperature, rainfall) from the end of April, a few days before we started detecting failed nests, and compared them with previous and subsequent years. In early May, when many pairs were incubating or caring for newly hatched chicks, we detected unusually low ambient temperatures and unusually high rainfall. This event likely caused that many first clutches failed completely, and that, compared with previous and subsequent years, fledging and breeding success was relatively low in those which were successful. There was also an increased proportion of replacement clutches that year. Anyway, the overall production of fledglings per breeding pair over the breeding season was 2.39, lower than other years. Although recruitment rate in 2003 was similar to other years, the lower production of fledglings in 2002 probably resulted in a decrease in the number of breeding pairs in the following years.

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.

Divergence in evolutionary potential of life history traits among wild populations is predicted by differences in climatic conditions.

Short-term adaptive evolution represents one of the primary mechanisms allowing species to persist in the face of global change. Predicting the adaptive response at the species level requires reliable estimates of the evolutionary potential of traits involved in adaptive responses, as well as understanding how evolutionary potential varies across a species' range. Theory suggests that spatial variation in the fitness landscape due to environmental variation will directly impact the evolutionary potential of traits. However, empirical evidence on the link between environmental variation and evolutionary potential across a species range in the wild is lacking. In this study, we estimate multivariate evolutionary potential (via the genetic variance-covariance matrix, or G-matrix) for six morphological and life history traits in 10 wild populations of great tits (Parus major) distributed across Europe. The G-matrix significantly varies in size, shape, and orientation across populations for both types of traits. For life history traits, the differences in G-matrix are larger when populations are more distant in their climatic niche. This suggests that local climates contribute to shaping the evolutionary potential of phenotypic traits that are strongly related to fitness. However, we found no difference in the overall evolutionary potential (i.e., G-matrix size) between populations closer to the core or the edge of the distribution area. This large-scale comparison of G-matrices across wild populations emphasizes that integrating variation in multivariate evolutionary potential is important to understand and predict species' adaptive responses to new selective pressures.

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.

Estimating Density Dependence, Environmental Variance, and Long-Term Selection on a Stage-Structured Life History.

AbstractA method for analyzing long-term demographic data on density-dependent stage-structured populations in a stochastic environment is derived to facilitate comparison of populations and species with different life histories. We assume that a weighted sum of stage abundances, N, exerts density dependence on stage-specific vital rates of survival and reproduction and that N has a small or moderate coefficient of variation. The dynamics of N are approximated as a univariate stochastic process governed by three key parameters: the density-independent growth rate, the net density dependence, and environmental variance in the life history. We show how to estimate the relative weighs of stages in N and the key parameters. Life history evolution represents a stochastic maximization of a simple function of the key parameters. The long-term selection gradient on the life history can be expressed as a vector of sensitivities of this function with respect to density-independent, density-dependent, and stochastic components of the vital rates. To illustrate the method, we analyze 38 years of demographic data on a great tit population, estimating the key parameters, which accurately predict the observed mean, coefficient of variation, and fluctuation rate of N; we also evaluate the long-term selection gradient on the life history.

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.

Global warming modifies the seasonal distribution of clutches on a Mediterranean great tit population

Global warming has multiple effects on phenological events on a wide range of plants and animals. Specifically, many bird species have advanced the start of their breeding season, which could also imply an extension in its duration and also a change in the distribution of clutches throughout the breeding season. We have tested whether this occurred in a population of Great Tits (Parus major), in Sagunto (eastern Spain). The increase of March temperatures between 1986 and 2019 was related to an advancement of the breeding season. Although the breeding season was longer in years with higher June temperatures, the length did not show a temporal trend throughout the study period. The clutches were more concentrated at the beginning of the season (increase in the skewness), while the kurtosis (“tailedness” of the distribution) or the modality did not change significantly. Finally, the number of “equally good months” for breeding (a combined measure of length and distribution) has not changed throughout the years. Overall, all these phenological changes probably caused the observed increased proportion of pairs laying two clutches per year. It remains to be studied whether this increase in reproductive effort has positive or negative impact on fitness.

Disentangling the causes of age-assortative mating in bird populations with contrasting life-history strategies.

Age shapes fundamental processes related to behaviour, survival and reproduction, where age influences reproductive success, non-random mating with respect to age can magnify or mitigate such effects. Consequently, the correlation in partners' age across a population may influence its productivity. Despite widespread evidence for age-assortative mating, little is known about what drives this assortment and its variation. Specifically, the relative importance of active (same-age mate preference) and passive processes (assortment as a consequence of other spatial or temporal effects) in driving age assortment is not well understood. In this paper, we compare breeding data from a great tit and mute swan population (51- and 31-year datasets, respectively) to tease apart the contributions of pair retention, cohort age structure and active age-related mate selection to age assortment in species with contrasting life histories. Both species show age-assortative mating and variable assortment between years. However, we demonstrate that the drivers of age assortment differ between the species, as expected from their life histories and resultant demographic differences. In great tits, pair fidelity has a weak effect on age-assortative mating through pair retention; variation in age assortment is primarily driven by fluctuations in age structure from variable juvenile recruitment. Age-assortative mating is, therefore, largely passive, with no evidence consistent with active age-related mate selection. In mute swans, age assortment is partly explained by pair retention, but not population age structure, and evidence exists for active age-assortative pairing. This difference is likely to result from shorter life-spans in great tits compared with mute swans, leading to fundamental differences in their population age structure, whereby a larger proportion of great tit populations consist of a single age cohort. In mute swans, age-assortative pairing through mate selection may also be driven by greater age-dependent variation in fitness. The study highlights the importance of considering how different life histories and demographic differences arising from these affect population processes that appear congruent across species. We suggest that future research should focus on uncovering the proximate mechanisms that lead to variation in active age-assortative mate selection (as seen in mute swans); and the consequences of variation in age structure on the ecological and social functioning of wild populations.

Parenting in the city: effects of urbanization on incubation behaviour and egg temperature in great tits, Parus major

With 68% of the world's population expected to live in cities by 2050, it is crucial to understand how animals respond to urbanization. Urban areas are associated with changes in microclimate, light, noise and human activity, which can affect animal physiology, behaviour and fitness. In particular, urbanization may affect how parents allocate time and energy between parental care and self-maintenance. In birds, a critical aspect of parental care is incubation, where parents must maintain egg temperatures for successful offspring development. However, incubation is energetically costly, and anthropogenic disturbances may affect parental decisions related to this cost. Determining whether urbanization affects incubation is crucial for understanding the differences between urban and nonurban populations in their reproductive performance and fitness. We measured incubation behaviour and egg temperature in one urban and one forest population of great tits, a uniparental incubator, using temperature loggers. We found that, compared to forest females, urban females spent more time incubating, took shorter off-bouts and ended their last daily off-bouts at a later hour. Furthermore, time spent off the nest, the number and duration of off-bouts and daily start of activity were related to ambient temperature; start of activity was related to clutch size; and time spent off the nest was related to the day of incubation. Despite urban females spending more time on the nest, eggs of both forest and urban females experienced relatively similar average egg temperatures, and urban eggs tended to experience greater variability in egg temperature. Furthermore, there were no differences between sites for any fitness-related offspring variable. Our results suggest that urbanization can influence the thermal environment of developing avian offspring by causing urban adults to be inefficient incubators through increased activity off the nest in the evening, differences in nest insulation and/or suboptimal incubation behaviour compared to forest counterparts. • Urbanization appears to influence parental incubation behaviour of great tits. • Urban birds spent more time warming their eggs compared to birds in the forest. • However, outcomes for offspring were similar. • Urban eggs and forest eggs experienced similar temperatures.

The effect of daylight exposure on the immune response and body condition of Great Tit nestlings

Natural light plays an important role in many aspects of birds’ biology, ecology and behaviour. Wild birds experience various light conditions, both in terms of its intensity and spectral properties, which vary within the nesting habitat of avian species. Recent study has shown that some bird species prefer to build their nests in bright nest sites. Light is one of the major factors affecting early development and condition of nestlings, but to date, this effect has not been tested in wild conditions. This study aims to answer whether birds could benefit from nesting in brighter light environment. Potential benefits may occur in nestlings’ condition, growth rate and breeding output. The research was carried out in the population of Great Tits (Parus major) breeding in two types nest boxes which differed with internal illumination. In “dark” boxes, the only source of light was the entrance hole. “Bright” boxes have additional side semi-transparent windows. We conducted two experiments in which light environment in nest box was set after (Experiment I) or before (Experiment II) birds settle in. Our study showed that light environment did not significantly affect nestlings body condition and growth. However, females in their first breeding attempt showed strong preference to the brightened nest boxes. In this group, the offspring were heavier and showed stronger immune responses than nestlings reared in dark nest boxes. Nestlings in bright nest boxes fledged about one day earlier. Nesting in brightened nest boxes may provide potential benefit for adult females in their first breeding attempt. The interaction between maternal effect and light environment is being discussed.

No relationship between chronotype and timing of breeding when variation in daily activity patterns across the breeding season is taken into account.

There is increasing evidence that individuals are consistent in the timing of their daily activities, and that individual variation in temporal behavior is related to the timing of reproduction. However, it remains unclear whether observed patterns relate to the timing of the onset of activity or whether an early onset of activity extends the time that is available for foraging. This may then again facilitate reproduction. Furthermore, the timing of activity onset and offset may vary across the breeding season, which may complicate studying the above‐mentioned relationships. Here, we examined in a wild population of great tits (Parus major) whether an early clutch initiation date may be related to an early onset of activity and/or to longer active daylengths. We also investigated how these parameters are affected by the date of measurement. To test these hypotheses, we measured emergence and entry time from/into the nest box as proxies for activity onset and offset in females during the egg laying phase. We then determined active daylength. Both emergence time and active daylength were related to clutch initiation date. However, a more detailed analysis showed that the timing of activities with respect to sunrise and sunset varied throughout the breeding season both within and among individuals. The observed positive relationships are hence potentially statistical artifacts. After methodologically correcting for this date effect, by using data from the pre‐egg laying phase, where all individuals were measured on the same days, neither of the relationships remained significant. Taking methodological pitfalls and temporal variation into account may hence be crucial for understanding the significance of chronotypes.

Prior experience of captivity affects behavioural responses to 'novel' environments.

Information ecology theory predicts that prior experience influences current behaviour, even if the information is acquired under a different context. However, when individuals are tested to quantify personality, cognition, or stress, we usually assume that the novelty of the test is consistent among individuals. Surprisingly, this ‘gambit of prior experience’ has rarely been explored. Therefore, here we make use of a wild population of great tits (Parus major) to test if prior experience of handling and captivity influences common measures of exploration (open field tests in two novel contexts: room and cage arenas), social response (simulated using a mirror), and behavioural stress (breathing rate). We found that birds with prior experience of captivity (caught previously for unrelated learning and foraging experiments) were more exploratory, but this depended on age: exploration and captivity experience (in terms of both absolute binary experience and the length of time spent in captivity) were associated more strongly in young (first-winter) birds than in adults. However, there was no association of prior experience of captivity with social response and breathing rate, and nor did the measures of exploration correlate. Together our results suggest that re-testing of individuals requires careful consideration, particularly for younger birds, and previous experiences can carry over and affect behaviours differently.

The scale of seasonal migrations in the urban part of the Great Tit population in the southeast of the West Siberian plain

Aim. This research aims to study the Great Tit breeding density, depending on the distance from preferred wintering sites with all other things being equal.Material and Methods. 36 route counts were carried out in 22 locations ranging up to 95 km away from the preferred wintering sites. The data was collected in the same weather and climatic zone. The route counts were carried out in the wintering season (January) and in the breeding season (16 April – 15 May) in 2014‐2018. To consider the influence of small settlements, where a smaller number of tits spends winter in comparison with a large city, sample sites were characterized by the population of people living within a radius of 25 km around the sample site.Results. Most Great Tits prefer to minimise the distance from their wintering sites to their nesting sites. The most preferred distance of seasonal movement is 10‐20 km from a large city, the main wintering place for this species.Conclusions. The density of the Great Tit population during the nesting period increases exponentially with the proximity of wintering sites. When more than 20 km from wintering sites, the density of the species during the nesting period does not change reliably. The maximum immanent density during the nesting period is achieved in places of favourable wintering of the species, that is, in a metropolis with agglomeration.

A longitudinal analysis of the growth rate and mass of tail feathers in a great tit population: ontogeny, genetic effects and relationship between both traits

Feathers have a diversity of functions in birds and are costly to produce, so their growth rate and mass can be reliable indicators of nutritional condition at the time of production. Despite the potential for feather metrics to advance our understanding of foraging, they are underused in avian ecology. One reason for this is the difficulty of interpreting whether individual variation is driven by ontogenetic, genetic or environmental effects, which is exacerbated by the fact that most analyses have been done on cross‐sectional data. We addressed this deficit using a longitudinal dataset of tail feathers collected from great titsParus majorto test for ontogenetic and genetic effects on growth rate, mass and length, while controlling for body/feather size differences and other confounding factors. First, we found that the type of moult episode and experimentally‐induced replacement differentially affected the length, mass and growth of feathers, providing evidence of an ontogenetic effect that should be considered when comparing these feather traits across individuals as a measure of condition. Second, we detected moderate to high repeatability and heritability values from parent–offspring regression for these three feather traits, which are suggestive of an underlying genetic component of variation. Third, we used a mean centring within‐individual approach to test whether feather growth rate and feather mass (length‐corrected) are indeed positively correlated with each other as overlapping indicators of body condition in birds, and found that this association, although positive, is weak and only significant between individuals. This suggests that both metrics are not so intimately linked as originally thought, and probably have different sensitivities to variation in foraging performance and ecological conditions. Together with the higher plasticity of feather growth rate compared to feather mass, our results support the idea that feather growth rate is better suited for examining short‐term responses to environmental variation.

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.