Variation in growth in the blue tit (Parus caeruleus)

The pattern and amount of phenotypic and genetic variation in growth was analysed using the infinite-dimensional model in natural populations of the blue tit (Parus caeruleus). A size-index was used to analyse patterns of variation and covariation across ages (2, 5, 8, 11 and 14 days). Genetic variation was analysed using mother-, father-, and midparent-offspring regressions in the blue tit. Phenotypic variation in growth was limited to a few dimensions only, while heritabilities were close to zero at all ages except at 2 days when a significant mother-offspring relation was found. The phenotypic variance-covariance matrix over ages was found to be singular, which suggests that there are trajectories for which there is no phenotypic variation, and which therefore they cannot evolve. The phenotypic growth trajectory associated with the largest amount of variation basically showed a high to moderately high covariation across ages. These results have macroevolutionary implications in terms of the outcome of selection which generally will be in terms of overall size changes but with a slow rate of change due to low heritabilities; an outcome consistent with macro-evolutionary patterns of morphological variation in birds. A simulation study supported the conclusion that high covariances among traits and ages can impose considerable constraint on a microevolutionary scale.     

Intersexual niche differentiation in the blue tit (Parus caeruleus)

The intersexual niche differentiation hypothesis, according to which sexual size dimorphism (SSD) has evolved to facilitate niche separation between the sexes, has received equivocal support in studies of birds. One explanation for the inconsistency of the results is that other factors crucial for reproductive success, such as variation in food abundance among territories, may obscure food niche differentiation between parents. We studied the effects of SSD, food availability and food demands of broods on niche differentiation in the blue tit ( Parus caeruleus ). Parents attending experimentally enlarged broods delivered the same amount, but more diverse food items, to their offspring than parents attending reduced broods, although this effect was significant in only one of the two study years. The time-lag (deviation from the peak food abundance in habitat in days) and squared time-lag were the most important determinants of the diversity of food items delivered to young and their condition (fitness surrogate). SSD, as measured by principal component scores of morphological traits, did not explain any variation neither in the diversity or amount of food provided, nor in offspring condition. These results suggest that synchronization of breeding with peak food availability is the most important determinant of diversity of food delivered to young and, more importantly, Their condition. No evidence for intersexual food niche differentiation was found; at best, it is likely to constitute only a weak selection pressure on the maintenance or enhancement of SSD in this system.     

Local specialization and maladaptation in the Mediterranean blue tit (Parus caeruleus)

Although the evolutionary interplay between gene flow and local adaptation of organisms in heterogeneous environments has been widely discussed from a theoretical point of view, few empirical studies have been designed to test predictions on the consequences of habitat patchiness on the evolution of life history traits. Using blue tits in Mediterranean habitat mosaics as a model, we defined two nested levels of habitat heterogeneity: an inter-regional level which compares two isolated landscapes (mainland, southern France vs the island of Corsica), and an intra-regional level which compares two habitat types within each landscape (deciduous vs evergreen trees). Deciduous habitats are more common than evergreen habitats on the mainland whereas the opposite is true on the island. Results suggest that: (1) on a regional scale, each population is specialized to the more common habitat, i.e. life history traits have evolved in such a way that breeding success is high; (2) in the less common habitats within each landscape, birds are clearly mistimed because they mismatch the best period of food availability, and hence their breeding success is lower; and (3) the density of the populations and the morphometry of the birds support the model of ideal despotic distribution. These results, which are supported by preliminary data on the genetic variation and gene flow of populations in the mainland landscape, are consistent with a source-sink model of population structure within each landscape. They are discussed in the framework of metapopulation theory and habitat selection models.     

Ultraviolet reflectance affects male-male interactions in the blue tit (Parus caeruleus ultramarinus)

Several animal species have been shown to use phenotypic traitsto assess the competitive ability of opponents and adjust theiraggressiveness depending on the likelihood to win the contest.In birds, these phenotypic traits usually involve patches ofcolored feathers. The benefit to harbor honest signals of malequality is the avoidance of wasteful aggressive interactions.Recent work has shown that ultraviolet (UV) plumage reflectanceis an important signal used by females during mate choice. Surprisingly,however, the role of UV signaling on intrasexual selection hasbeen neglected. In the present study, we aimed to test whetherUV reflectance of crown feathers was used as a signal of malecompetitive ability during male-male interactions. Breedingmale blue tits (Parus caeruleus ultramarinus) were exposed duringthe female egg-laying period to blue tit taxidermic mounts witheither control or reduced UV reflectance of crown feathers.In agreement with the prediction that UV reflectance advertisesmale quality, we found that breeding blue tits behaved lessaggressively toward the UV-reduced decoy. To our knowledge,this is the first experimental evidence suggesting a role forUV signaling on intrasexual selection.     

Egg quality and parental ornamentation in the blue tit Parus caeruleus

Components of avian egg quality often show remarkable variation between females and yet the causes of variation in this fitness-related trait remain poorly understood. We investigated the egg size and yolk carotenoid investment of blue tit Parus caeruleus females in relation to multiple parental traits. We show that females produce eggs with more colourful yolks when mated to ultraviolet (UV) attractive males. Since yolk carotenoids may enhance offspring fitness, but may be available to females in limiting amounts, the correlation we found suggests adaptive egg quality adjustment in response to mate attractiveness, but an experimental approach is required to rule out alternative explanations. Yolk colour was also correlated with laying date and yolk mass. Though the effect of laying date could not be explained by ambient temperature during egg formation, it suggests a proximate constraint of general carotenoid availability on yolk composition. Egg size was not affected by male attractiveness; however, females with a brighter crown produced larger eggs, suggesting that crown coloration might indicate individual quality in females too.     

Different population structures in northern and southern populations of the European blue tit (Parus caeruleus)

We sequenced the control region of the mitochondrial DNA from a sample of six European blue tit populations to investigate the phylogeography of Parus species. Along a transect from Barcelona, Spain to Oulu, Finland, the blue tit showed a different phylogeographic structure than the great tit and the willow tit. The southernmost sample from Barcelona consisted of two widely divergent maternal lineages (nucleotide divergence, π = 0.30%), a situation also found earlier in the French Alps. The more northern populations had a relatively uniform structure (π = 0.19%) with distinctive marks of a growing population, thus resembling the great tit populations (π = 0.19%). The amount of genetic variation among blue tits is lower than in the willow tit (π = 0.53%). This probably reflects a smaller long-term effective population size in the great tit and the blue tit than in the willow tit. The different genetic structure of the Barcelona population vs. the rest had an influence on the estimated population parameters, which are calculated based on the assumptions of genetic equilibrium of the populations.     

Glacial history and colonization of Europe by the blue tit Parus caeruleus

Mitochondrial control region sequences from European populations of the blue tit Parus caeruleus were used to reveal the Pleistocene history and the post-glacial recolonization of Europe by the species. The southern subspecies, P. c. ogliastrae was found to represent a stable population with isolation-by-distance structure harboring a lot of genetic variation, and the northern subspecies P. c. caeruleus a recently bottlenecked and expanded population. We suggest that after the last Ice Ages, the subspecies have colonized Europe from two different southern refuges following previously proposed general recolonization routes from the Balkans to northern and Central Europe, and from the Iberian Peninsula north- and eastwards. The two subspecies form a wide secondary contact zone extending from southern Spain to southern France.     

Natural selection on immune responsiveness in blue tits Parus caeruleus

Abstract.— What is the form of natural selection on immune responsiveness? For a population at evolutionary equilibrium, there are two different scenarios. First, it is generally assumed that immune defense has both benefits and costs. If variation in immune responsiveness is due to variation in how individuals trade off these costs and benefits, one would expect immune responsiveness to be subject to stabilizing selection. Second, it is well known that an individual's immune responsiveness is often dependent on its overall condition. If immune responsiveness is condition‐dependent, one would expect immune responsiveness to be under positive directional selection. We would therefore expect that the form of natural selection on immune responsiveness depends on the relative magnitude of these two sources of variation: variation in how individuals trade off the costs and benefits of defense, and variation in condition. We measured primary and secondary antibody responsiveness to diphtheria‐tetanus vaccine in blue tits during winter and investigated the relationship between responsiveness and survival to the following breeding season. We use responsiveness to these antigens as measures of an individual's ability or propensity to mount an antibody response in case of an infection. Interestingly, different measures of responsiveness were subject to different selective regimes: primary responsiveness to diphtheria was subject to stabilizing selection, whereas secondary responsiveness to tetanus was subject to positive directional selection. In contrast, there was no significant selection on primary responsiveness to tetanus or secondary responsiveness to diphtheria. The finding of stabilizing selection on a measure of responsiveness is evidence that immune defense can incur fitness costs; a central but little‐tested assumption of theories of the ecology and evolution of immunological defense. The finding of directional selection on a measure of responsiveness is consistent with the idea that immune responsiveness is condition‐dependent, although we cannot rule out the alternative explanation that the population is not at evolutionary equilibrium with respect to this trait.     

Two blue tit Parus caeruleus populations from Corsica differ in social dominance

Although the causes and consequences of social dominance have been examined extensively, avian studies have rarely focused on between-population differences in social dominance. On the island of Corsica, two resident blue tit Parus caeruleus populations 25 km apart differ significantly in body size measures, timing and effort of reproduction, and song structure, and some of these population differences have a genetic basis. Because earlier avian studies have shown that social dominance is influenced by body size or mass, we predicted that individuals from these two blue tit populations would also differ in their ability to dominate other individuals. Consistent with this prediction, we found that male blue tits of these two populations differ in social dominance, and that heavier or larger individuals dominate lighter or smaller ones in aviary experiments. We propose that social dominance may serve to maintain phenotypic population differentiation at a micro-geographic scale by acting as a barrier to dispersal.     

Unusual mitochondrial DNA polymorphism in two local populations of blue tit Parus caeruleus

Mitochondrial DNA (mtDNA) from 25 blue tits Parus caeruleus sampled from two populations of the Grenoble region (France) was assayed for polymorphism with 17 restriction endonucleases. Nine genotypes were found. Several mtDNA genotypes were also analysed by amplification via the polymerase chain reaction (PCR) and direct sequencing of 903 bp of the cytochrome b gene. The mtDNA polymorphism is greater in P. caeruleus than in other comparable bird species and results from the presence of two clearly differentiated mitochondrial lineages. Using the data of restriction polymorphism, the mean sequence divergence between individuals of the two lineages is 1.23%. Therefore, P. caeruleus should fall into the category II of phylogeographic pattern sensu Avise et al. (1987): discontinuous mtDNA genotypes which co-occur in the same region. P. caeruleus, like humans and other mobile species with high gene flow, seems to have lost its geographic structure in terms of mtDNA phylogeny. This unusual mitochondrial polymorphism can be explained by the recent admixture of two long-term isolated populations. This could be accounted for by two different scenarios. One assumes a simultaneous post-glacial colonization of the Grenoble region by two isolated European populations of P. caeruleus. Alternatively, hybridization between P. caeruleus and P. cyanus could have caused the observed pattern of mtDNA variation.     

Identification of hypervariable single locus minisatellite DNA probes in the blue tit Parus caeruleus

We report the isolation of a set of hypervariable minisatellite DNA sequences from a blue tit Parus caeruleus genomic DNA library. In our strategy, we cloned a minisatellite-rich DNA fraction into a charomid vector. The resulting cosmid library was screened with the two minisatellite DNA probes 33.6 and 33.15 for recombinants containing a minisatellite DNA insert. A total of 233 positive clones were isolated. Of 37 clones that have been analysed, nine gave polymorphic signals and can be used as single locus probes (SLPs). Four of the SLPs were investigated in more detail. The number of alleles, the heterozygosity and the mutation rate were estimated. Linkage analysis revealed that two of these loci were linked. The SLPs are of value to studies of the mating system and reproductive success in the blue tit, and may also be useful in population genetic studies.     

No effect of parental quality or extrapair paternity on brood sex ratio in the blue tit (Parus caeruleus)

Sex allocation theory predicts that parents should manipulatebrood sex ratio in order to maximise the combined reproductivevalue of their progeny. Females mating with high quality malesshould, therefore, be expected to produce brood sex ratiosbiased towards sons, as male offspring would receive a relativelygreater advantage from inheritance of their father's characteristicsthan would their female siblings. Furthermore, it has been suggested that sex allocation in chicks fathered through extrapair fertilizations should also be biased towards sons. Contraryto these predictions, we found no evidence that the distributionof sex ratios in a sample of 1483 chicks from 154 broods ofblue tits (Parus caeruleus) deviated significantly from thatof a binomial distribution around an even sex ratio. In addition,we found no significant effect on brood sex ratio of the individualquality of either parent as indicated by their biometrics, feather mite loads, time of breeding, or parental survival. This suggeststhat females in our population were either unable to manipulateoffspring sex allocation or did not do so because selectionpressures were not strong enough to produce a significant shiftaway from random sex allocation. The paternity of 986 chicks from 103 broods was determined using DNA microsatellite typing.Extrapair males sired 115 chicks (11.7%) from 41 broods (39.8%).There was no significant effect of paternity (within-pair versusextrapair) on the sex of individual offspring. We suggest that,in addition to the weakness of selection pressures, the possiblemechanisms responsible for the allocation of sex may not besufficiently accurate to control offspring sex at the levelof the individual egg.     

Environmental factors shape cloacal bacterial assemblages in great tit Parus major and blue tit P. caeruleus nestlings

Despite their potential ecological and evolutionary importance, factors shaping the composition of bacterial communities in wild vertebrate populations remain poorly understood. The goal of this study was to examine the relative contributions of environmental factors and genetic factors (e.g. species and common origin) to the variation of cloacal bacterial assemblages in wild bird nestlings. We conducted a partial cross-fostering experiment with two passerine species, the great tit Parus major and the blue tit P. caeruleus, sharing similar habitats and breeding biology. Nestlings of the two species were exchanged four days after hatching and cloacal bacteria were sampled nine days later. The structure of cloacal bacterial communities was determined by Ribosomal Intergenic Spacer Analysis. Our results showed that each nestling displayed a unique bacterial community. Furthermore, nestlings raised in the same nest shared significantly similar bacterial communities. The similarity of bacterial community was higher among heterospecific siblings raised within the same nest than between biological siblings raised in separate nests. Effects of common origin between species could not be detected and, if present, were dominated by nest-based short-term environmental effects. Our results show that growth conditions within nests and individually based endogenous factors have significant effects on cloacal bacteria assemblages and could affect post-fledging condition.     

Extrapair paternity in the blue tit (Parus caeruleus) : female choice, male charateristics, and offspring quality

Extrapair paternity is common in many birds, and it is now generallyaccepted that female choice plays an important role. However,die benefits that females obtain from extrapair paternity aremuch less dear. To test the hypothesis that females obtain indirectfitness benefits, we studied paternity in a blue tit populationover 4 years. Extrapair paternity occurred in 31-47% of allnests and accounted for 11-14% of all offspring. Most malesthat fathered extrapair young did not lose paternity themselves,males never "exchanged" paternity, and within nests the extrapairoffspring were usually fathered by a single male. Comparisonsbetween males that did and did not lose paternity and pairwisecomparisons between the extrapair male(s) and the within-pairmale showed that successful males had longer tarsi and sangon average longer strophes during the dawn chorus. Successfulmales weighed less (relative to their size) during the nettlingstage, but nevertheless they survived better. Male age did notinfluence their likelihood of losing paternity, but extrapairmales were usually older than the within-pair male they cuckolded.Within nests with mixed paternity, extrapair young were morelikely to survive than within-pair young in cases of partialbrood mortality. Our data also suggest that extrapair offspringwere more likely to be males. Because extrapair males were usuallyclose neighbors, male quality should be considered relativeto the quality of the neighbors. Despite this, we found consistencyin female choice over years. Our observations provide supportfor the hypothesis that female blue tits engage in extrapaircopulations to obtain good genes for their offspring.     

Genetic variation and causes of genotype-environment interaction in the body size of blue tit (Parus caeruleus).

In several studies of natural populations of birds, the heritability of body size estimated by parent-offspring regression has been lower when offspring have developed in poor feeding regimens than when they developed in good feeding regimens. This has led to the suggestion that adaptation under poor regimens may be constrained by lack of genetic variation. We examined the influence of environmental conditions on expression of genetic variation in body size of nestling blue tits (Parus caeruleus) by raising full sibs in artificially reduced and enlarged broods, corresponding to good and poor feeding regimens, respectively. Individuals grown in the poor regimen attained smaller body size than their sibs grown in the good regimen. However, there was among-family variation in response to the treatments--i.e., genotype-environment interactions (GEIs). Partitioning the GEI variance into contributions attributable to (1) differences in the among-family genetic variance between the treatments and (2) imperfect correlation of genotypic values across treatments identified the latter as the main cause of the GEI. Parent-offspring regressions were not significantly different when offspring were reared in the good environment (h2 = 0.75) vs. when they were reared in the poor environment (h2 = 0.63). Thus, there was little evidence that genetic variance in body size was lower under the poor conditions than under the good conditions. These results do not support the view that the genetic potential for adaptation to poor feeding conditions is less than that for adaptation to good conditions, but they do suggest that different genotypes may be favored under the different conditions.     

Separating the effects of predation risk and interrupted foraging upon mass changes in the blue tit Parus caeruleus

The optimal amount of reserves that a small bird should carry depends upon a number of factors, including the availability of food and environmental predation risk levels. Theory predicts that, if predation risk increases, then a bird should maintain a lower level of reserves. Previous experiments have given mixed results: some have shown reduced reserves and some, increased reserves. However, the birds in these studies may have been interpreting a staged predation event as a period when they were unable to feed rather than a change in predation risk: theory predicts that, if the food supply within the environment is variable, then reserves should be increased. In the present study, we presented blue tits (Parus caeruleus) with a potential predator and compared this response (which could have been potentially confounded by perceived interruption effects) with a response to an actual interruption in the environment during both long and short daytime lengths. During long (but not short) days, the birds responded in line with theoretical predictions by increasing their reserves in response to interruption and reducing them in response to predation. These results are examined in the light of other experimental manipulations and we discuss how well experimental tests have tested the predictions made by theoretical models.     

Male-biased sex ratio among unhatched eggs in great tit Parus major, blue tit P. caeruleus and collared flycatcher Ficedula albicollis

An increasing number of studies have demonstrated that brood sex ratios are frequently unequal, but the proximate mechanisms underlying this deviation are largely unknown. In the current study we analysed deviation from expected 1:1 sex ratio among dead embryos from unhatched eggs collected from partially unhatched clutches of three passerine bird species. We showed that male embryos were significantly overrepresented among unhatched eggs of great tit Parus major , blue tit P. caeruleus and collared flycatcher Ficedula albicollis . Moreover, the bias in sex ratio differed among species and tended to differ among study years. We discuss several hypotheses to explain the observed male bias among unhatched eggs. We conclude that sex specific embryo mortality may contribute to explain the observed variation in sex ratios in several species of wild birds and that sexing unhatched eggs is important in studies of sex ratio allocation.     

Experimentally increased reproductive effort alters telomere length in the blue tit (Cyanistes caeruleus)

Telomeres have recently been suggested to play important role in ageing and are considered to be a reliable ageing biomarkers. The life history theory predicts that costs of reproduction should be expressed in terms of accelerated senescence, and some empirical studies do confirm such presumption. Thus, a link between reproductive effort and telomere dynamics should be anticipated. Recent studies have indeed demonstrated that reproduction may trigger telomere loss, but actual impact of reproductive effort has not received adequate attention in experimental studies. Here, we experimentally manipulated reproductive effort by increasing the brood size in the wild blue tit (Cyanistes caeruleus). We show that parents attending enlarged broods experienced larger yearly telomere decay in comparison to control birds attending unaltered broods. In addition, we demonstrate that the change in telomere length differs between sexes, but this effect was independent from our treatment. To our knowledge, this is the first experimental study in the wild revealing that telomere dynamics may be linked to reproductive effort. Thus, telomere shortening may constitute one of the potential proximate mechanisms mediating the costs of reproduction.     

Genetic and environmental components of growth in nestling blue tits (Parus caeruleus)

We investigated the effect of brood‐size mediated food availability on the genetic and environmental components of nestling growth in the blue tit (Parus caeruleus), using a cross‐fostering technique. We found genetic variation for body size at most nestling ages, and for duration of mass increase, but not of tarsus growth. Hence, nestling growth in our study population seems to have the potential to evolve further. Furthermore, significant genotype–environment interactions indicated heritable variation in reaction norms of growth rates and growth periods, i.e. that our study population had a heritable plasticity in the growth response to environmental conditions. The decreasing phenotypic variance with nestling age indicated compensatory growth in all body traits. Furthermore, the period of weight increase was longer for nestlings growing up in enlarged broods, while there was no difference to reduced broods in the period of tarsus growth. At fledging, birds in enlarged broods had shorter tarsi and lower weights than birds in reduced broods, but there was no difference in wing length or body condition between the two experimental groups. The observed flexibility in nestling growth suggests that growing nestlings are able to respond adaptively to food constraint by protecting the growth of ecologically important traits.