Sympatric divergence and clinal variation in multiple coloration traits of Ficedula flycatchers.

Geographic variation in phenotypes plays a key role in fundamental evolutionary processes such as local adaptation, population differentiation and speciation, but the selective forces behind it are rarely known. We found support for the hypothesis that geographic variation in plumage traits of the pied flycatcher Ficedula hypoleuca is explained by character displacement with the collared flycatcher Ficedula albicollis in the contact zone. The plumage traits of the pied flycatcher differed strongly from the more conspicuous collared flycatcher in a sympatric area but increased in conspicuousness with increasing distance to there. Phenotypic differentiation (PST ) was higher than that in neutral genetic markers (FST ), and the effect of geographic distance remained when statistically controlling for neutral genetic differentiation. This suggests that a cline created by character displacement and gene flow explains phenotypic variation across the distribution of this species. The different plumage traits of the pied flycatcher are strongly to moderately correlated, indicating that they evolve non-independently from each other. The flycatchers provide an example of plumage patterns diverging in two species that differ in several aspects of appearance. The divergence in sympatry and convergence in allopatry in these birds provide a possibility to study the evolutionary mechanisms behind the highly divergent avian plumage patterns.

Size differentiation in Finnish house sparrows follows Bergmann's rule with evidence of local adaptation.

Bergmann's rule predicts that individuals are larger in more poleward populations and that this size gradient has an adaptive basis. Hence, phenotypic divergence in size traits between populations (PST ) is expected to exceed the level of divergence by drift alone (FST ). We measured 16 skeletal traits, body mass and wing length in 409 male and 296 female house sparrows Passer domesticus sampled in 12 populations throughout Finland, where the species has its northernmost European distributional margin. Morphometric differentiation across populations (PST ) was compared with differentiation in 13 microsatellites (FST ). We find that twelve traits phenotypically diverged more than FST in both sexes, and an additional two traits diverged in males. The phenotypic divergence exceeded FST in several traits to such a degree that findings were robust also to strong between-population environmental effects. Divergence was particularly strong in dimensions of the bill, making it a strong candidate for the study of adaptive molecular genetic divergence. Divergent traits increased in size in more northern populations. We conclude that house sparrows show evidence of an adaptive latitudinal size gradient consistent with Bergmann's rule on the modest spatial scale of ca. 600 km.

Exploring patterns of variation in clutch size-density reaction norms in a wild passerine bird.

Negative density dependence of clutch size is a ubiquitous characteristic of avian populations and is partly due to within-individual phenotypic plasticity. Yet, very little is known about the extent to which individuals differ in their degree of phenotypic plasticity, whether such variation has a genetic basis and whether level of plasticity can thus evolve in response to selection. Using 18 years of data of a Dutch great tit population (Parus major), we show that females reduced clutch size with increasing population density (slopes of the reaction norms), differed strongly in their average clutch size (elevations of the reaction norms) at the population-mean density and that the latter variation was partly heritable. In contrast, we could not detect individual variation in phenotypic plasticity ('I × E'). Level of plasticity is thus not likely to evolve in response to selection in this population. Observed clutch sizes deviated more from the estimated individual reaction norms in certain years and densities, implying that the within-individual between-year variance (so-called residual variance) of clutch size was heterogeneous with respect to these factors. Given the observational nature of this study, experimental manipulation of density is now warranted to confirm the causality of the observed density effects. Our analyses demonstrate that failure to acknowledge this heterogeneity would have inflated the estimate of 'I × E' and led to misinterpretation of the data. This paper thereby emphasizes the fact that heterogeneity in residuals can provide biologically insightful information about the ecological processes underlying the data.

Quantitative genetics of behavioural reaction norms: genetic correlations between personality and behavioural plasticity vary across stickleback populations.

Behavioural ecologists have proposed various evolutionary mechanisms as to why different personality types coexist. Our ability to understand the evolutionary trajectories of personality traits requires insights from the quantitative genetics of behavioural reaction norms. We assayed > 1000 pedigreed stickleback for initial exploration behaviour of a novel environment, and subsequent changes in exploration over a few hours, representing their capacity to adjust their behaviour to changes in perceived novelty and risk. We found heritable variation in both the average level of exploration and behavioural plasticity, and population differences in the sign of the genetic correlation between these two reaction norm components. The phenotypic correlation was not a good indicator of the genetic correlation, implying that quantitative genetics are necessary to appropriately evaluate evolutionary hypotheses in cases such as these. Our findings therefore have important implications for future studies concerning the evolution of personality and plasticity.

Whither Pst? The approximation of Qst by Pst in evolutionary and conservation biology.

Local adaptation through natural selection can be inferred in case the additive genetic divergence in a quantitative trait across populations (Q(st)) exceeds the neutral expectation based on differentiation of neutral alleles across these populations (e.g. F(st)). As such, measuring Q(st) in relation to neutral differentiation presents a first-line investigation applicable in evolutionary biology (selection on functional genes) and conservation biology (identification of locally adapted coding genes). However, many species, especially those in need of conservation actions, are not amenable for the kind of breeding design required to estimate either narrow- or broad-sense Q(st). In such cases, Q(st) has been approximated by the phenotypic divergence in a trait across populations (P(st)). I here argue that the critical aspect for how well P(st) approximates Q(st) depends on the extent that additive genetic effects determine variation between populations relative to within populations. I review how the sensitivity of conclusions regarding local adaptation based on P(st) have been evaluated in the literature and find that many studies make a anticonservative null assumption in estimating P(st) and/or use a nonconservative approach to explore sensitivity of their conclusions. Data from two studies that have provided a second, independent assessment of selection in their system suggest that P(st)-F(st) comparisons should be interpreted very conservatively. I conclude with recommendations for improving the robustness of the inferences drawn from comparing P(st) with neutral differentiation.

Blood parasites mediate morph-specific maintenance costs in a colour polymorphic wild bird.

Parasites can mediate profound negative effects on host fitness. Colour polymorphism has been suggested to covary genetically with intrinsic physiological properties. Tawny owl colour polymorphism is highly heritable with two main morphs, grey and brown. We show that experimental medication acts to reduce blood parasites and that medicated grey females maintain body mass during breeding, whereas medicated brown females decline in body mass similar to control females of both morphs. We find no effect of medication on general immunoglobulin levels, antigen-specific humoral response or H/L ratio. In the descriptive data, both morphs have similar blood parasite infection rates, but blood parasite infection is associated with decreased body mass in brown but not in grey females. We conclude that blood parasite infection primarily has somatic costs, which differ between the two highly heritable tawny owl colour morphs with more pronounced costs in the grey (little pigmented) morph than in the brown (heavily pigmented) morph. Because our descriptive results imply the opposite pattern, our findings highlight the need of experimental manipulation when studying heritable variation in hosts' response to parasitism.

Low genetic differentiation in a sedentary bird: house sparrow population genetics in a contiguous landscape.

The house sparrow Passer domesticus has been declining in abundance in many localities, including Finland. We studied the genetic diversity and differentiation of the house sparrow populations across Finland in the 1980s, at the onset of the species' decline in abundance. We genotyped 472 adult males (the less dispersive sex) from 13 locations in Finland (covering a range of 400 × 800 km) and one in Sweden (Stockholm) for 13 polymorphic microsatellite markers. Our analysis of Finnish ringing records showed that natal dispersal distances are limited (90% <16 km), which confirmed earlier finding from other countries. The Finnish populations were panmictic, and genetically very homogeneous and the limited dispersal was sufficiently large to maintain their connectivity. However, all Finnish populations differed significantly from the Stockholm population, even though direct geographical distance to it was often smaller than among Finnish populations. Hence, the open sea between Finland and Sweden appears to form a dispersal barrier for this species, whereas dispersal is much less constrained across the Finnish mainland (which lacks geographical barriers). Our findings provide a benchmark for conservation biologists and emphasize the influence of landscape structure on gene flow.

The rate of ageing in a long-lived bird is not heritable.

A senescent decline in performance occurs in late age in many organisms, and is thought to be partly due to additive genetic effects. Here annual fitness, estimated as the age-specific sum of survival and reproduction, was used to test for genetic variance in ageing in a population of common gulls, Larus canus. Data on 3986 individuals collected over a 34-year period indicate a dramatic senescent decline in late life. We also find that annual fitness is heritable and that individuals vary in their rates of ageing. However, counter to theoretical expectations, we find no support for a heritable component to the variance in rates of senescence. Increases in the among-individual (permanent environment) and residual variance components initiate an increase in the total phenotypic variance for annual fitness with age. This finding suggests that older birds are more sensitive to environmental effects, and that old age causes an overall pattern of declining h(2) of annual fitness. Our findings suggest that individual-specific factors do have a role in determining the rate of senescence in this population, but that additive genetic variance for the rate of senescence is either absent or small.

Heritability, plasticity and canalization of Ural owl egg size in a cyclic environment.

Avian egg size is highly variable on the population level, but is considered inflexible on the individual level. On the basis of 2969 measurements of individual eggs collected during 1981-2005, we analysed heritability, plasticity and selection on egg size in the Ural owl, a long-lived bird that preys on voles. Vole abundance varied in a 3-year cycle, creating varying food supply across the cycle's phases. Ural owl egg size is heritable (h(2) = 60%). Ural owls lay larger eggs in improved food conditions. On the basis of repeated breeding records of 59 females that bred in all vole cycle phases, we show that intra-individual adjustment (plasticity) explained 22.4% of the variation in egg size across phases. Egg size was under stabilizing selection. Extremely small and extremely large eggs had reduced hatchability, and individuals who laid either large or small eggs had lower lifetime fledgling production than the ones laying intermediately sized eggs. Our findings illustrate how maternal investment in egg size can both be heritable and highly responsive to variable environmental conditions, and suggest that variation in the investment in egg size across individuals is canalized.

A possible link between parasite defence and residual reproduction.

Life-history theory centres around trade-offs between current and future reproduction, but we have little understanding of how such trade-offs are mediated. We supplementary fed Ural owls (Strix uralensis) during the nestling period and quantified parents' current and future life-history components as well as their physiological health by monitoring haematocrit, leucocyte profile, intra- and extracellular blood parasites. Feeding led to reduced parental effort but did not improve offspring viability, male parasite defence, or parental survival. Intracellular leucocytozoan infection was reduced in fed females which lasted to the following year's reproductive season (carry-over effect), when fed females also laid larger and earlier clutches. Leucocytozoon infection therefore may mediate the life-history trade-off between current and residual reproduction in this species.

The evolutionary ecology of individual phenotypic plasticity in wild populations.

The ability of individual organisms to alter morphological and life-history traits in response to the conditions they experience is an example of phenotypic plasticity which is fundamental to any population's ability to deal with short-term environmental change. We currently know little about the prevalence, and evolutionary and ecological causes and consequences of variation in life history plasticity in the wild. Here we outline an analytical framework, utilizing the reaction norm concept and random regression statistical models, to assess the between-individual variation in life history plasticity that may underlie population level responses to the environment at both phenotypic and genetic levels. We discuss applications of this framework to date in wild vertebrate populations, and illustrate how natural selection and ecological constraint may alter a population's response to the environment through their effects at the individual level. Finally, we present future directions and challenges for research into individual plasticity.

The evolution of fitness in life-history theory.

Theory concerning the evolution of life history (the schedule of reproduction and survival) focuses on describing the life history which maximises fitness. Although there is an intuitive link between life history and fitness, there are in fact several measures of the 'black box' concept of fitness. There has been a debate in the bio-mathematical literature on the predictive difference between the two most commonly used measures; intrinsic rate of increase r and net reproductive ratio R0. Although both measures aim to describe fitness, models using one of the measures may predict the opposite of similar models using the other measure, which is clearly undesirable. Here, I review the evolution of these fitness measures over the last four decades, the predictive differences between these measures and the resulting shift of the fitness concept. I focus in particular on some recent developments, which have solved the dilemma of predictive differences between these fitness measures by explicitly acknowledging the game-theoretical nature of life-history evolution.