Can dominance genetic variance be ignored in evolutionary quantitative genetic analyses of wild populations?

Accurately estimating genetic variance components is important for studying evolution in the wild. Empirical work on domesticated and wild outbred populations suggests that dominance genetic variance represents a substantial part of genetic variance, and theoretical work predicts that ignoring dominance can inflate estimates of additive genetic variance. Whether this issue is pervasive in natural systems is unknown, because we lack estimates of dominance variance in wild populations obtained in situ. Here, we estimate dominance and additive genetic variance, maternal variance, and other sources of nongenetic variance in eight traits measured in over 9000 wild nestlings linked through a genetically resolved pedigree. We find that dominance variance, when estimable, does not statistically differ from zero and represents a modest amount (2-36%) of genetic variance. Simulations show that (1) inferences of all variance components for an average trait are unbiased; (2) the power to detect dominance variance is low; (3) ignoring dominance can mildly inflate additive genetic variance and heritability estimates but such inflation becomes substantial when maternal effects are also ignored. These findings hence suggest that dominance is a small source of phenotypic variance in the wild and highlight the importance of proper model construction for accurately estimating evolutionary potential.     

Latitudinal variation in blue tit and great tit nest characteristics indicates environmental adjustment

Aim The laying of eggs and the building of a nest structure to accommodate them are two of the defining characteristics of members of the class Aves. Nest structures vary considerably across avian taxa and for many species the structure of the completed nest can have important consequences both for parents and their offspring. While nest characteristics are expected to vary adaptively in response to environmental conditions, large‐scale spatial variation in nest characteristics has been largely overlooked. Here, we examine the effects of latitudinal variation in spring temperatures on nest characteristics, including insulatory properties, and reproductive success of blue tits, Cyanistes caeruleus, and great tits, Parus major. Location Great Britain. Methods Nests and reproductive data were collected from seven study sites, spread over 5° of latitude. The nest insulatory properties were then determined before the nests were separated into nest base material and cup lining material. Results As spring temperatures increased with decreasing latitude, the mass of the nest base material did not vary in either species, while the mass of the cup lining material and nest insulatory properties decreased in both species. This suggests that in response to increasing temperatures the breeding female reduces the mass of the cup lining material, thereby maintaining an appropriate microclimate for incubating and brooding. The mean first egg date of both species advanced with decreasing latitude and increasing spring temperatures, although clutch size and brood size at hatching and fledging did not vary. Main conclusions This is the first study to demonstrate that the nest‐construction behaviour of birds varies in response to large‐scale spatial variation in ambient temperatures. Therefore, nest composition reliably indicates environmental conditions and we suggest that studies of nest structure may be sentinels for the early signs of rapid climate change.     

Heritability of resting metabolic rate in a wild population of blue tits

We report the first study with the aim to estimate heritability in a wild population, a nest box breeding population of blue tits. We estimated heritability as well as genetic and phenotypic correlations of resting metabolic rate (RMR), body mass and tarsus length with an animal model based on data from a split cross‐fostering experiment with brood size manipulations. RMR and body mass, but not tarsus length, showed significant levels of explained variation but for different underlying reasons. In body mass, the contribution to the explained variation is mainly because of a strong brood effect, while in RMR it is mainly because of a high heritability. The additive variance in RMR was significant and the heritability was estimated to 0.59. The estimates of heritability of body mass (0.08) and tarsus length (0.00) were both low and based on nonsignificant additive variances. Thus, given the low heritability (and additive variances) in body mass and tarsus length the potential for direct selection on RMR independent of the two traits is high in this population. However, the strong phenotypic correlation between RMR and mass (0.643 ± 0.079) was partly accounted for by a potentially strong, although highly uncertain, genetic correlation (1.178 ± 0.456) between the two traits. This indicates that the additive variance of body mass, although low, might still somewhat constrain the independent evolvability of RMR.     

Manipulation of male attractiveness induces rapid changes in avian maternal yolk androgen deposition

Avian eggs contain maternal androgens that may adjust offspringdevelopment to environmental conditions. We review evidenceand functional explanations for the relationship between androgenconcentrations in avian eggs and male attractiveness. Experimentalstudies in captive birds show generally positive relationships,but results from correlational and experimental field studiesare less consistent, perhaps because they lack a within-femaledesign to control for confounding between-female variation.We analyzed the effect of male attractiveness on yolk levelsof maternal androgens in a wild bird, using a correlationaland experimental approach with a within-female design. We manipulatedthe sexually selected UV coloration of the crown feathers ofmale blue tits (Cyanistes caeruleus) after their female hadlaid the second egg and measured the subsequent effect on androgenconcentrations (testosterone and androstenedione) in the fifth,seventh, and ninth eggs relative to that in the second egg.Levels of testosterone, but not androstenedione, in eggs 5 and7 were higher for control (attractive) than for UV-reduced (unattractive)males. This effect disappeared in the ninth egg, coincidingwith the recovery of UV coloration after manipulation. Thissuggests that females are capable of rapid adjustments of testosteronedeposition in response to changes in their mate's ornamentalplumage. However, androgen concentrations in the second eggand pretreatment male crown coloration were not correlated.Possibly, the combination of relatively small variation in UVcoloration before treatment and the influence of unknown confoundingvariables in the correlative approach resulted in insufficientstatistical power to detect such a correlation.     

Evolutionary quantitative genetics of behavioral responses to handling in a wild passerine

Behavioral differences between individuals that are consistent over time characterize animal personality. The existence of such consistency contrasts to the expectation based on classical behavioral theory that facultative behavior maximizes individual fitness. Here, we study two personality traits (aggression and breath rate during handling) in a wild population of blue tits during 2007–2012. Handling aggression and breath rate were moderately heritable (h² = 0.35 and 0.20, respectively) and not genetically correlated (r_A = 0.06) in adult blue tits, which permits them to evolve independently. Reciprocal cross‐fostering (2007–2010) showed that offspring reared by more aggressive males have a higher probability to recruit. In addition, offspring reared by pairs mated assortatively for handling aggression had a higher recruitment probability, which is the first evidence that both parents' personalities influence their reproductive success in the wild in a manner independent of their genetic effects. Handling aggression was not subjected to survival selection in either sex, but slow‐breathing females had a higher annual probability of survival as revealed by capture–mark–recapture analysis. We find no evidence for temporal fluctuations in selection, and thus conclude that directional selection (via different fitness components) acts on these two heritable personality traits. Our findings show that blue tit personality has predictable fitness consequences, but that facultative adjustment of an individual's personality to match the fitness maximum is likely constrained by the genetic architecture of personality. In the face of directional selection, the presence of heritable variation in personality suggests the existence of a trade‐off that we have not identified yet.     

Wild fledgling tits do not mob in response to conspecific or heterospecific mobbing calls

Mobbing, where birds harass a predator through a combination of vocalizations and stereotyped behaviours, is an effective anti-predator behaviour for many species. Mobbing may be particularly important for juveniles, as these individuals are often more vulnerable than adults. Although the component behaviours of mobbing are often considered to be un-learned, there are few confirmatory data, and the developmental trajectory of mobbing is unknown. In this study, we tested whether conspecific or heterospecific mobbing calls initiated mobbing behaviour in juvenile Blue Tits Cyanistes caeruleus. We located wild adult and recently fledged juvenile Blue Tits and presented them with playback recordings of adult conspecific (Blue Tit) and heterospecific (Great Tit Parus major) mobbing alarm calls. Although adult birds readily mob in response to these types of playbacks, juveniles did not exhibit characteristic mobbing behaviour. Some juveniles did, however, exhibit individual components of mobbing behaviour found in mobbing, despite not producing adult-like mobbing behaviour in response to either conspecific or heterospecific playback. These results suggest that, although birds might be capable of mobbing as juveniles, the associations between the non-vocal stereotyped mobbing behaviours and mobbing calls may be learned.     

Impaired flight ability--a cost of reproduction in female blue tits

When prey are attacked by predators, escape ability has an obvious influence on the probability of survival. Laboratory studieshave suggested that flight performance of female birds mightbe affected by egg production. This is the first study of changesin take-off ability, and thus potentially in predation risk,during reproduction in wild birds. We trapped individual maleand female blue tits repeatedly during the breeding season.Females were 14% heavier and flew 20% slower (probably as aconsequence of a lower ratio of flight muscle to body mass)during the egg-laying period than after the eggs had hatched.However, flight muscle size did not change to compensate for changes in body mass over this period. In contrast, males showedno changes in either body mass, muscle size, or flight abilityover the same period. Furthermore, the impairment of flightin females increased with the proportion of the clutch thathad been laid, an effect that was independent of body mass and muscle size. This indicates that egg production causes additional physiological changes in the female body that produce impairedlocomotor performance. We suggest that courtship feeding offemale blue tits by their mates might reduce predation riskduring the period when female take-off ability is impairedby reducing the time females have to spend foraging and thusreducing the time they are exposed to increased predation.     

Climate change and breeding parameters of great and blue tits throughout the western Palaearctic

Increasing evidence suggests that climate change has consequences on avian breeding phenology. Here, variations in laying date and clutch size of great tit Parus major and blue tit Parus caeruleus within and between breeding populations through the western Palaearctic are examined in relation to climatic fluctuations, measured by the winter North Atlantic Oscillation (NAO) index. Within and across breeding sites, laying date was related to winter‐NAO index such that great and blue tit females lay earlier after warmer, moister winters (positive values of winter NAO‐index). The present study shows that for most populations there is an advancement of laying date, but the rate of change with respect to NAO significantly differed geographically across the western Palaearctic and did not differ between species. However, clutch size of great and blue tits was not affected by climatic fluctuations, presumably because the whole season is being shifted, but not in relation to food supplies. These combined analyses for the two species controlled for potentially confounding variables such as latitude, longitude, elevation and habitat of each study site.     

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.     

Determinants of distribution and prevalence of avian malaria in blue tit populations across Europe: separating host and parasite effects

Although avian malarial parasites are globally distributed, the factors that affect the geographical distribution and local prevalence of different parasite lineages across host populations or species are still poorly understood. Based on the intense screening of avian malarial parasites in nine European blue tit populations, we studied whether distribution ranges as well as local adaptation, host specialization and phylogenetic relationships can determine the observed prevalences within populations. We found that prevalence differed consistently between parasite lineages and host populations, indicating that the transmission success of parasites is lineage specific but is partly shaped by locality-specific effects. We also found that the lineage-specific estimate of prevalence was related to the distribution range of parasites: lineages found in more host populations were generally more prevalent within these populations. Additionally, parasites with high prevalence that were also widely distributed among blue tit populations were also found to infect more host species. These findings suggest that parasites reaching high local prevalence can also realize wide distribution at a global scale that can have further consequences for host specialization. Although phylogenetic relationships among parasites did not predict prevalence, we detected a close match between a tree based on the geographic distance of the host populations and the parasite phylogenetic tree, implying that neighbouring host populations shared a related parasite fauna.