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.     

COMPARATIVE METHODS AT THE SPECIES LEVEL: GEOGRAPHIC VARIATION IN MORPHOLOGY AND GROUP SIZE IN GREY-CROWNED BABBLERS (POMATOSTOMUS TEMPORALIS)

We show that a new comparative method that sheds light on evolutionary trends among species may also illuminate trends within species. This finding comes from a phylogenetic autocorrelation analysis of morphological traits among individuals sampled from ten populations of a cooperatively breeding songbird, the Grey-crowned Babbler (Pomatostomus temporalis). Highly variable mitochondrial DNA (mtDNA) sequences from both the eastern (Pomatostomus temporalis temporalis) and western (Pomatostomus temporalis rubeculus) lineages were used to define genetic distances among 120 individuals and to estimate correlations among individuals in wing length, tarsus length, and body weight via an intraspecific weighting matrix. The autoregressive model effectively removed intraspecific correlations for all three morphological variables, and the proportion of the total phenotypic variance due to genealogical relationships varied from 0.68 (weight) to 0.23 (tarsus). The analysis revealed correlations among the specific components of traits, in which none were previously detected (type-I error) and diminished correlations that appeared strong when phylogeny was ignored. Group size was the only trait for which the autoregressive model failed to remove intraspecific correlations, a result likely due to the plasticity, convergence, and clinal variation in this trait in both the eastern and western lineages. The autocorrelation analysis weakened significant negative correlations between group size and total values for wing length and body weight, but the interpretation of this result depends on the adaptive significance ascribed to the “phylogenetic component” of trait values removed by the analysis. Comparative methods employing distance matrices are one useful way of summarizing the pattern of nonhierarchical relationships among conspecific individuals (“tokogenetic” relationships, sensu Hennig).     

Does larder-hoarding affect the condition of chicks in urban kestrels?

Hoarding by urban kestrels and its subsequent exploitation significantly increases the frequency and regularity of feeding during the early stage of the nestling period. Twelve kestrels’ nests situated in various part of the city were chosen to test whether hoarding could affect the chicks’ condition. Four nests (with 22 nestlings) were provided with food — one dead mouse per chick per day (F group) and 8 (43 nestlings) were used as a control group (NF). Nestlings were weighed daily, and the length of their tarsus was also measured when the oldest chick in the nest was 11 days old. 50% of the NF nests experienced a decrease in the number of nestlings but no nestlings were lost in any of the F nests. However, no significant differences were found between the groups either in body mass or tarsus length. A possible reason was the behaviour of the adults which could be interpreted as the regulation of the frequency of prey delivery depending on its presence or absence in the nest.     

树麻雀表型的环境适应进化

树麻雀(Passer montanus)分布范围广、海拔梯度大,也是人类活动的伴随物种。对中国837个样本的10个形态特征与温度、日照、海拔和风速等4个主要环境因子进行相关分析,结果显示:树麻雀的体重、嘴裂、翅长、尾长、跗跖长、脑骨宽、眼间距与日照因子显著相关(P0.05),体重、体长、翅长、尾长、跗跖长与海拔因子显著相关(P0.05),体重、嘴峰、翅长、脑骨长与温度因子显著相关(P0.05),表明树麻雀的形态指标易随环境因子的变化而变化。通过控制经度和海拔两个变量,对形态指标与纬度的偏相关分析表明,体重、翅长、脑骨长和脑骨宽与纬度呈显著正相关(P0.05),体表突出部分嘴峰、嘴裂与纬度呈显著负相关(P0.05),即随着纬度的升高,树麻雀身体逐渐变大,符合贝格曼规律;体表突出部分嘴峰和嘴裂随纬度升高变短,符合阿伦规律。飞行能力与海拔因子呈极显著正相关(n=92,r=0.217,P=0.038),表明树麻雀在高海拔地区具有更强的飞行能力,这也许是它成为广布种的重要原因。     

Genetic variance in temperature dependent adult size deriving from physiological genetic variation at temperature boundaries

An increase in genetic variation in body size has often been observed under stress; an increase in dominance variance and interaction variance as well as in additive genetic variance has been reported. The increase in genetic variation must be caused by physiological mechanisms that are specific to adverse environments. A model is proposed to explain the occurrence of an increase in genetic variation in body size in Drosophila at extreme temperatures. The model has parameters specific to the low- and high-temperature regions of the viable range. Additive genetic variation in the boundary temperatures leads to a marked increase in additive genetic variation in development rate and body size at extreme temperatures. Additive genetic variation in the temperature sensitivity in the low- and high-temperature regions adds non-additive genetic variation. Development rate shows patterns in additive genetic variation that differ from the patterns of genetic variation in body size; therefore, the genetic correlation between development rate and body size changes sign repeatedly as a function of temperature. The existence of dominance in the genetic variation in the boundary temperatures or in the low- and high-temperature sensitivities leads to a higher total genetic variance due to higher dominance and interaction variance, for both development rate and body size. This revised version was published online in July 2006 with corrections to the Cover Date.     

HERITABILITY AND SELECTION ON TARSUS LENGTH IN THE PIED FLYCATCHER (FICEDULA HYPOLEUCA)

The heritability estimate (±SE) for tarsus length in the pied flycatcher is 0.53 ± 0.10, based on mother-offspring regressions. The heritability is almost the same (0.50 ± 0.22) for offspring transferred to other nests and reared by foster parents, whereas there is no resemblance (0.04 ± 0.23) between the offspring and their foster mothers. Hence, the nest environment does not affect parent-offspring resemblance. However, a full-sib correlation yields an estimate of the heritability twice as high as the parent-offspring regression did, indicating that shared environment effects, which are not correlated with the tarsus length of parents, must be important. An environmental deviation due to food factors affecting tarsus length is demonstrated. The most important food factors are probably associated with 1) polygyny, which leads to reduced paternal feeding at secondary nests, 2) high breeding density, and 3) progress of the breeding season. All three are associated with reductions in offspring mean tarsus length. We estimate selection on tarsus length for the major components of lifetime reproductive success. Offspring with the shortest tarsi have reduced survival from fledging until breeding, and males with tarsus length close to the mean are most successful in attracting mates. Clutch size increases with female tarsus length, except for individuals with very long tarsi. In general, directional selection is weak, but stabilizing selection is rather strong for survival and male mating success.     

Effects of temperature and nest heat exposure on nestling growth,dehydration and survival in a Mediterranean hole‐nesting passerine

Variable environments impose constraints on adaptation by modifying selection gradients unpredictably. Optimal bird development requires an adequate thermal range, outside which temperatures can alter nestling physiology, condition and survival. We studied the effect of temperature and nest heat exposure on the reproductive success of a population of double‐brooded Spotless Starlings Sturnus unicolor breeding in a nestbox colony in central Spain with a marked intra‐seasonal variation in temperature. We assessed whether the effect of temperature differed between first and second broods, thus constraining optimal nest‐site choice. Ambient temperature changed greatly during the chick‐rearing period and had a strong influence on nestling mass and all body size measures we recorded, although patterns of clutch size or nestling mortality were not influenced. This effect differed between first and second broods: nestlings were found to have longer wings and bills with increasing temperature in first broods, whereas the effect was the opposite in second broods. Ambient temperature was not related to nestling body mass or tarsus‐length in first broods, but in second broods, nestlings were lighter and had smaller tarsi with higher ambient temperatures. The exposure of nestboxes to heat influenced nestling morphology: heat exposure index was negatively related to nestling body mass and wing‐length in second broods, but not in first broods. Furthermore, there was a positive relationship between nest heat exposure and nestling dehydration. Our results suggest that optimal nest choice is constrained by varying environmental conditions in birds breeding over prolonged periods, and that there should be selection for parents to switch from sun‐exposed to sun‐protected nest‐sites as the season progresses. However, nest‐site availability and competition for sites are likely to impose constraints on this choice.     

Sexual size dimorphism and positive assortative mating in red-backed shrike Lanius collurio: an adaptive value?

During field studies in 1997–1999 in South Bohemia (Czech Republic), we found significant differences in size between the sexes in a local breeding population of red-backed shrike Lanius collurio. Males were significantly larger than females for wing length and tarsus length, but had smaller body mass than females. However, there was considerable overlap in the ranges of these parameters between the sexes. Interestingly, pairs were formed at random with respect to wing length and tarsus length, but assortative mating was significant for body mass/body condition. Among tested variables, only male wing length correlated significantly with nestling body mass at day 7. However, clutch size and the number of fledglings strongly depended on differences in tarsus length between mates, but not on body size of mates. Individual improvements in foraging skills and/or courtship feeding rates are proposed as possible explanations for these findings.     

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.     

Natural selection on body size traits in a long‐lived bird,the snow petrel Pagodroma nivea

I consider the possibility of selection favouring large body size in a population of snow petrels (Pagodroma nivea), a long‐lived seabird species. I measured natural selection on body size traits in a population from 1987 to 1998. There was evidence of selection on body size associated with fecundity and survival. Directional selection on bill length and stabilizing selection on tarsus length associated with reproductive success were detected among males. Selection associated with survival favoured males with longer bills. However, selection was weak in all cases. No evidence of selection acting on female body size traits was detected. Offspring–parents regression suggested that bill length and tarsus length were heritable. Although I was able to identify the targets of selection in this population, I could not demonstrate the ecological implications of both tarsus length and bill length variation. The selection on male, but not on female, body size traits suggests factors such as intrasexual competition for nests and/or mates rather than factors such as feeding efficiency as mechanisms of selection on bill size.