Sexual reproduction is inherently interactive, especially in animal species such as humans that exhibit extended pair bonding. Yet we have little knowledge of the role of male characteristics and their evolutionary impact on reproductive behavioural phenotypes, to the extent that biologists typically consider component traits (e.g. reproductive timing) as female-specific. Based on extensive genealogical data detailing the life-histories of 6,435 human mothers born across four centuries of modern history, we use an animal modelling approach to estimate the indirect genetic effect of men on the reproductive phenotype of their partners. These analyses show that a woman’s reproductive timing (age at first birth) is influenced by her partner’s genotype. This indirect genetic effect is positively correlated with the direct genetic effect expressed in women, such that total heritable variance in this trait is doubled when heritable partner effects are considered. Our study thus suggests that much of the heritable variation in women’s reproductive timing is mediated via partner effects, and that the evolutionary potential of this trait is far greater than previously appreciated.

In natural populations, quantitative trait dynamics often do not appear to follow evolutionary predictions. Despite abundant examples of natural selection acting on heritable traits, conclusive evidence for contemporary adaptive evolution remains rare for wild vertebrate populations, and phenotypic stasis seems to be the norm. This so-called "stasis paradox" highlights our inability to predict evolutionary change, which is especially concerning within the context of rapid anthropogenic environmental change. While the causes underlying the stasis paradox are hotly debated, comprehensive attempts aiming at a resolution are lacking. Here, we apply a quantitative genetic framework to individual-based long-term data for a wild rodent population and show that despite a positive association between body mass and fitness, there has been a genetic change towards lower body mass. The latter represents an adaptive response to viability selection favouring juveniles growing up to become relatively small adults, i.e. with a low potential adult mass, which presumably complete their development earlier. This selection is particularly strong towards the end of the snow-free season, and it has intensified in recent years, coinciding which a change in snowfall patterns. Importantly, neither the negative evolutionary change, nor the selective pressures that drive it, are apparent on the phenotypic level, where they are masked by phenotypic plasticity and a non causal (i.e. non genetic) positive association between body mass and fitness, respectively. Estimating selection at the genetic level enabled us to uncover adaptive evolution in action and to identify the corresponding phenotypic selective pressure. We thereby demonstrate that natural populations can show a rapid and adaptive evolutionary response to a novel selective pressure, and that explicitly (quantitative) genetic models are able to provide us with an understanding of the causes and consequences of selection that is superior to purely phenotypic estimates of selection and evolutionary change.

Biologists are increasingly interested in decomposing trait dynamics into underlying processes, such as evolution, plasticity and demography. Four important frameworks that allow for such a decomposition are the quantitative genetic animal model (AM), the ‘Geber’ method (GM), the age-structured Price equation (APE) and the integral projection model (IPM). However, as these frameworks have largely been developed independently, they differ in the assumptions they make, the data they require, as well as their outcomes and interpretation. Here, we evaluate how each framework decomposes trait dynamics into underlying processes. To do so, we apply them to simulated data for a hypothetical animal population. Individual body size was affected by, among others, genes, maternal effects and food intake. We simulated scenarios with and without selection on body size and with high and low heritability. The APE and IPM provided similar results, as did the AM and GM, with important differences between the former and the latter. All frameworks detected positive contributions of selection in the high but not in the low selection scenarios. However, only the AM and GM distinguished between the high and low heritability scenarios. Furthermore, the AM and GM revealed a high contribution of plasticity. The APE and IPM attributed most of the change in body size to ontogenetic growth and inheritance, where the latter captures the combined effects of plasticity, maternal effects and heritability. We show how these apparent discrepancies are mostly due to differences in aims and definitions. For example, the APE and IPM capture selection, whereas the AM and GM focus on the response to selection. Furthermore, the frameworks differ in the processes that are ascribed to plasticity and in how they take into account demography. We conclude that no single framework provides the ‘true’ contributions of evolution, plasticity and demography. Instead, different research questions require different frameworks. A thorough understanding of the different definitions of their components is necessary for selecting the most appropriate framework for the question at hand and for making biologically meaningful inferences. This work thus supports both future analysis and the careful interpretation of existing work.

Inspection of the data that accompany Pruitt and Krauel's study of individual vari- ation in satiation threshold and a comparison of these data with the Materials and Methods and Results sections of the paper have revealed a number of issues that cast doubts on the reliability of the data and any results based on these data. In particular, we show that, following our analyses, the data are unlikely to have been obtained using the study design outlined in the publication and that statistical analyses of these data provide results that differ in important ways from those reported. These findings illustrate the importance of making raw data and analysis code available for the rigour and reproducibility of the scientific literature.

Resting metabolic rate (RMR) is a potentially important axis of physiological adaptation to the thermal environment. However, our understanding of the causes and consequences of individual variation in RMR in the wild is hampered by a lack of data, as well as analytical challenges. RMR measurements in the wild are generally characterized by large measurement errors and a strong dependency on mass. The latter is problematic when assessing the ability of RMR to evolve independently of mass. Mixed models provide a powerful and flexible tool to tackle these challenges, but they have rarely been used to estimate repeatability of mass-independent RMR from field data. We used respirometry to obtain repeated measurements of RMR in a long-term study population of snow voles (Chionomys nivalis) inhabiting an environment subject to large circadian and seasonal fluctuations in temperature. Using both uni- and bivariate mixed models, we quantify individual repeatability in RMR and decompose repeatability into mass-dependent and mass-independent components, while accounting for measurement error. RMR varies among individuals, i.e. is repeatable (R=0.46), and strongly co-varies with BM. Indeed, much of the repeatability of RMR is attributable to individual variation in BM, and the repeatability of mass-independent RMR is reduced by 41% to R=0.27. These empirical results suggest that the evolutionary potential of RMR independent of mass may be severely constrained. This study illustrates how to leverage bivariate mixed models to model field data for metabolic traits, correct for measurement error, and decompose the relative importance of mass-dependent and mass- independent physiological variation.

AbstractEvolutionary adaptation through genetic change requires genetic variation and is a key mechanism enabling species to persist in changing environments. Although a substantial body of work has focused on understanding how and why additive genetic variance (VA) differs among traits within species, we still know little about how they vary among species. Here we make a first attempt at testing for interspecific variation in two complementary measures of VA and the role of phylogeny in shaping this variation. To this end, we performed a phylogenetic comparative analysis using 1822 narrow‐sense heritability (h2) for 68 species of birds and mammals and 378 coefficients of additive genetic variance (CVA) estimates for 23 species. Controlling for within‐species variation attributable to estimation method and trait type, we found some interspecific variation in h2 (~15%) but not CVA. Although suggestive of interspecific variation in the importance of non‐(additive) genetic sources of variance, sample sizes were insufficient to test this hypothesis directly. Additionally, although power was low, no phylogenetic signal was detected for either measure. Hence, while this suggests interspecific variation in VA is probably small, our understanding of interspecific variation in the adaptive potential of wild vertebrate populations is currently hampered by data limitations, a scarcity of CVA estimates and a measure of their uncertainty in particular.

An individual's lifetime reproductive success (LRS) measures its realized genetic contributions to the next generation, but how well does it predict this over longer periods? Here we use human genealogical data to estimate expected individual genetic contributions (IGC) and quantify the degree to which LRS, relative to other fitness proxies, predicts IGC over longer periods. This allows an identification of the life-history stages that are most important in shaping variation in IGC. We use historical genealogical data from two non-isolated local populations in Switzerland to estimate the stabilized IGC for 2230 individuals approximately 10 generations after they were born. We find that LRS explains 30% less variation in IGC than the best predictor of IGC, the number of grandoffspring. However, albeit less precise than the number of grandoffspring, we show that LRS does provide an unbiased prediction of IGC. Furthermore, it predicts IGC better than lifespan, and accounting for offspring survival to adulthood does not improve the explanatory power. Overall, our findings demonstrate the value of human genealogical data to evolutionary biology and suggest that reproduction—more than lifespan or offspring survival—impacts the long-term genetic contributions of historic humans, even in a population with appreciable migration.

The rate of adaptive evolution, the contribution of selection to genetic changes that increase mean fitness, is determined by the additive genetic variance in individual relative fitness. To date, there are few robust estimates of this parameter for natural populations, and it is therefore unclear whether adaptive evolution can play a meaningful role in short-term population dynamics. We developed and applied quantitative genetic methods to long-term datasets from 19 wild bird and mammal populations and found that, while estimates vary between populations, additive genetic variance in relative fitness is often substantial and, on average, twice that of previous estimates. We show that these rates of contemporary adaptive evolution can affect population dynamics and hence that natural selection has the potential to partly mitigate effects of current environmental change.

Body height and body mass index (BMI) are associated with later life outcomes in present and historical populations. We examine the case study of the Swiss Alpine canton of Glarus, which was highly industrialised at the beginning of the 20th century. Our study links conscription registers to genealogical registers at the individual level in Switzerland for the first time. We analyse whether body height, BMI, socioeconomic position (HISCLASS), region of residence, fitness to serve (as a proxy for health status in a military context), and goitre status (as a proxy for iodine deficiency) in young adulthood are associated with lifespan. We transcribed conscription records of 1073 men born between 1905 and 1907 and recruited between 1925 and 1927 (coverage birth cohorts 96%). of the 827 young men residing within the canton, we were able to identify 635 (76.8%) in the cantonal genealogical register. Using body height, chest circumference, and upper arm circumference, we estimated BMI. We find socioeconomic differences for height and estimated BMI at conscription age. Young men with a recorded goitre were taller. We also present a positive association between body height and lifespan, with small men being particularly disadvantaged. In a small subsample of two municipalities, we estimated the heritability of height to be 65%.

AbstractAldabrachelys gigantea (Aldabra giant tortoise) is one of only two giant tortoise species left in the world and survives as a single wild population of over 100,000 individuals on Aldabra Atoll, Seychelles. Despite this large current population size, the species faces an uncertain future because of its extremely restricted distribution range and high vulnerability to the projected consequences of climate change. Captive‐bred A. gigantea are increasingly used in rewilding programs across the region, where they are introduced to replace extinct giant tortoises in an attempt to functionally resurrect degraded island ecosystems. However, there has been little consideration of the current levels of genetic variation and differentiation within and among the islands on Aldabra. As previous microsatellite studies were inconclusive, we combined low‐coverage and double‐digest restriction‐associated DNA (ddRAD) sequencing to analyze samples from 33 tortoises (11 from each main island). Using 5426 variant sites within the tortoise genome, we detected patterns of within‐island population structure, but no differentiation between the islands. These unexpected results highlight the importance of using genome‐wide genetic markers to capture higher‐resolution genetic structure to inform future management plans, even in a seemingly panmictic population. We show that low‐coverage ddRAD sequencing provides an affordable alternative approach to conservation genomic projects of non‐model species with large genomes.

Historically, mothers producing twins gave birth, on average, more often than non-twinners. This observation has been interpreted as twinners having higher intrinsic fertility - a tendency to conceive easily irrespective of age and other factors - which has shaped both hypotheses about why twinning persists and varies across populations, and the design of medical studies on female fertility. Here we show in >20k pre-industrial European mothers that this interpretation results from an ecological fallacy: twinners had more births not due to higher intrinsic fertility, but because mothers that gave birth more accumulated more opportunities to produce twins. Controlling for variation in the exposure to the risk of twinning reveals that mothers with higher twinning propensity - a physiological predisposition to producing twins - had fewer births, and when twin mortality was high, fewer offspring reaching adulthood. Twinning rates may thus be driven by variation in its mortality costs, rather than variation in intrinsic fertility.

This thesis is in two distinct parts. Part 1 focusses on the optimization of provisioning strategies and Part 2 on the evolution of morphological castes.

(1) in the Hymenoptera, the transition from mass to progressive provisioning, where mothers feed offspring gradually as they grow, is associated with sociality. Because progressively provisioning mothers usually feed more than one offspring simultaneously, they have to make decisions about which offspring to feed next. The genus Ammophila is especially tractable as a study system because different offspring are in spatially separate nests, so that maternal activities can be clearly associated with particular offspring. We use an individual based simulation in combination with field data from A. pubescens to investigate how provisioning strategies might be optimized. We find provisioning the oldest offspring first significantly reduces the risk of both conspecific and heterospecific parasitism and outperforms our other strategies. We also find that this strategy could have evolved without the use of memory to keep track of the relative ages of offspring, despite incurring heavy sampling costs through parasitism. We then go on to consider the impacts of poor weather on informing optimal provisioning strategies.

(2) the evolution of eusociality is regarded as one of the major evolutionary transitions (METs), where units that previously reproduced independently function as a single, more complex entity or group. Conflict and control are particularly important in understanding METs, since minimising within-group conflicts is critical for efficient functioning of the larger entity. Eusocial societies, characterised by a division of labour between worker and queen castes, represent one of the most studied METs. In advanced eusocial societies with morphological castes, queens and provisioners often influence the morphology of workers through a variety of mechanisms, thus potentially constraining worker behaviour. For morphology to constrain behaviour, offspring must then modify their behaviour in response to morphology, but the correlation between behaviour and morphology in reality might not be perfect. We explore some of the conditions under which morphological castes may arise and the factors constraining their evolution. We find that a mother’s certainty about her offsprings’ behaviour, including the ability for offspring to modify their behaviour in response to their morphology, are key in determining when we expect morphological castes to evolve.

ABSTRACTAn individual’s lifetime reproductive success (LRS) measures its realised genetic contributions to the next generation, but how well does it predict these over longer periods? Here we use human genealogical data to estimate expected individual genetic contributions (IGC) and quantify the degree to which LRS, relative to other fitness proxies, predicts IGC over longer periods in natural populations. This allows an identification of the life-history stages that are most important in shaping variation in IGC. We use historical genealogical data from two non-isolated local populations in Switzerland to estimate the stabilised IGC for 2,230 individuals ~10 generations after they were born. We find that LRS explains 30% less variation in IGC than the best predictor of IGC, the number of grandoffspring. However, albeit less precise than the number of grandoffspring, we show that LRS does provide an unbiased prediction of IGC and overall predicts IGC better than lifespan and similarly when accounting for offspring survival to adulthood. Overall, our findings demonstrate the value of human genealogy data to evolutionary biology and showing that reproduction - more than lifespan or offspring survival - impacts the long-term genetic contributions of historic humans, even in a population with appreciable migration.

AbstractPremiseMaternal effects have been demonstrated to affect offspring performance in many organisms, and in plants, seeds are important mediators of these effects. Some woody plant species maintain long‐lasting canopy seed banks as an adaptation to wildfires. Importantly, these seeds stored in serotinous cones are produced by the mother plant under varying ontogenetic and physiological conditions.MethodsWe sampled the canopy seed bank of a highly serotinous population of Pinus pinaster to test whether maternal age and growth and the environmental conditions during each crop year affected seed mass and ultimately germination and early survival. After determining retrospectively the year of each seed cohort, we followed germination and early survival in a semi‐natural common garden.ResultsSeed mass was related to maternal age and growth at the time of seed production; i.e. slow‐growing, older mothers had smaller seeds, and fast‐growing, young mothers had larger seeds, which could be interpreted either as a proxy of senescence or as a maternal strategy. Seed mass had a positive effect on germination success, but aside from differences in seed mass, maternal age had a negative effect and diameter had a positive effect on germination timing and subsequent survival.ConclusionsThe results highlight the importance of maternal conditions combined with seed mass in shaping seedling establishment. Our findings open new insights in the offspring performance deriving from long‐term canopy seed banks, which may have high relevance for plant adaptation.

Begging for food, a conspicuous solicitation display, is common in a variety of taxa, and it has received extensive research attention in a parent-offspring context. Both theoretical models and empirical evidence suggest that offspring begging can be an honest signal of hunger or a mediator of competition between siblings. At a behavioural mechanistic level, begging for food can be a form of harassment aimed at persuading those in possession of food to share. Food sharing, defined as the transfer of a defendable food item from one individual to another, can vary considerably between species, age-classes and food type and abundance. We investigated the determinants of begging and food-sharing behaviours in Asian small-clawed otters (Aonyx cinereus), a group-living species that commonly exhibits begging in captivity. We presented two captive otter populations with three food types that varied in exploitation complexity, in three different abundances. We predicted that begging rates would be highest when food was in lowest abundance and hardest to exploit, and that increased begging would lead to increased food sharing. We found that, over time, increased begging rates were indeed correlated with increased food transfers, but neither food type complexity nor abundance affected begging or sharing rates. However, age category was significantly associated with begging and food sharing rates: juvenile otters begged more and shared less than adult otters. The results from this first experimental study on begging and food sharing within the Mustelid family begin to reveal some of the drivers of these behaviours.

SUMMARYThe evolutionary potential of organisms depends on the presence of sufficient genetic variation for traits subject to selection, as well as on the genetic covariances among them. While genetic variation ultimately derives from mutation, theory predicts the depletion of genetic (co)variation under consistent directional or stabilizing selection in natural populations. We estimated and compared additive genetic (co)variances for several standard life history traits, including some for which this has never been assessed, before and after 24 generations of artificial selection on male size in the yellow dung flyScathophaga stercoraria(Diptera: Scathophagidae) using a series of standard half-sib breeding experiments. As predicted, genetic variances (VA), heritabilities (h2) and evolvabilities (IA) of body size, development time, first clutch size, and female age at first clutch were lower after selection. As independent selection lines were crossed prior to testing, we can rule out that this reduction is due to genetic drift. In contrast to the variances, and against expectation, the additive genetic correlations between the sexes for development time and body size remained strong and positive (rA =0.8–0.9), while the genetic correlation between these traits within the sexes tended to strengthen (but not significantly so). Our study documents that the effect of selection on genetic variance is predictable, whereas that on genetic correlations is not.

Sexual reproduction is inherently interactive, especially in animal species such as humans that exhibit extended pair bonding. Yet we have little knowledge of the role of male characteristics and their evolutionary impact on reproductive behavioural phenotypes, to the extent that biologists typically consider component traits (e.g. reproductive timing) as female-specific. Based on extensive genealogical data detailing the life-histories of 6,435 human mothers born across four centuries of modern history, we use an animal modelling approach to estimate the indirect genetic effect of men on the reproductive phenotype of their partners. These analyses show that a woman’s reproductive timing (age at first birth) is influenced by her partner’s genotype. This indirect genetic effect is positively correlated with the direct genetic effect expressed in women, such that total heritable variance in this trait is doubled when heritable partner effects are considered. Our study thus suggests that much of the heritable variation in women’s reproductive timing is mediated via partner effects, and that the evolutionary potential of this trait is far greater than previously appreciated.

In natural populations, quantitative trait dynamics often do not appear to follow evolutionary predictions. Despite abundant examples of natural selection acting on heritable traits, conclusive evidence for contemporary adaptive evolution remains rare for wild vertebrate populations, and phenotypic stasis seems to be the norm. This so-called "stasis paradox" highlights our inability to predict evolutionary change, which is especially concerning within the context of rapid anthropogenic environmental change. While the causes underlying the stasis paradox are hotly debated, comprehensive attempts aiming at a resolution are lacking. Here, we apply a quantitative genetic framework to individual-based long-term data for a wild rodent population and show that despite a positive association between body mass and fitness, there has been a genetic change towards lower body mass. The latter represents an adaptive response to viability selection favouring juveniles growing up to become relatively small adults, i.e. with a low potential adult mass, which presumably complete their development earlier. This selection is particularly strong towards the end of the snow-free season, and it has intensified in recent years, coinciding which a change in snowfall patterns. Importantly, neither the negative evolutionary change, nor the selective pressures that drive it, are apparent on the phenotypic level, where they are masked by phenotypic plasticity and a non causal (i.e. non genetic) positive association between body mass and fitness, respectively. Estimating selection at the genetic level enabled us to uncover adaptive evolution in action and to identify the corresponding phenotypic selective pressure. We thereby demonstrate that natural populations can show a rapid and adaptive evolutionary response to a novel selective pressure, and that explicitly (quantitative) genetic models are able to provide us with an understanding of the causes and consequences of selection that is superior to purely phenotypic estimates of selection and evolutionary change.

. Studies on the evolution of cooperative behaviour are typically confined to understanding its adaptive value. It is equally essential, however, to understand its potential to evolve, requiring knowledge about the phenotypic consistency and genetic basis of cooperative behaviour. While previous observational studies reported considerably high heritabilities of helping behaviour in cooperatively breeding vertebrates, experimental studies disentangling the relevant genetic and non-genetic components of cooperative behaviour are lacking. In a half-sibling breeding experiment, we investigated the repeatability and heritability of three major helping behaviours performed by subordinates of the cooperatively breeding fish
. Neolamprologus pulcher
. To experimentally manipulate the amount of help needed in a territory, we raised the fish in two environments differing in egg predation risk. All three helping behaviours were significantly repeatable, but had very low heritabilities. The high within-individual consistencies were predominantly due to maternal and permanent environment effects. The perceived egg predation risk had no effect on helping, but social interactions significantly influenced helping propensities. Our results reveal that developmentally plastic adjustments of provided help to social context shape cooperative phenotypes, whereas heritable genetic variation plays a minor role.
.

Biologists are increasingly interested in decomposing trait dynamics into underlying processes, such as evolution, plasticity and demography. Four important frameworks that allow for such a decomposition are the quantitative genetic animal model (AM), the ‘Geber’ method (GM), the age-structured Price equation (APE) and the integral projection model (IPM). However, as these frameworks have largely been developed independently, they differ in the assumptions they make, the data they require, as well as their outcomes and interpretation. Here, we evaluate how each framework decomposes trait dynamics into underlying processes. To do so, we apply them to simulated data for a hypothetical animal population. Individual body size was affected by, among others, genes, maternal effects and food intake. We simulated scenarios with and without selection on body size and with high and low heritability. The APE and IPM provided similar results, as did the AM and GM, with important differences between the former and the latter. All frameworks detected positive contributions of selection in the high but not in the low selection scenarios. However, only the AM and GM distinguished between the high and low heritability scenarios. Furthermore, the AM and GM revealed a high contribution of plasticity. The APE and IPM attributed most of the change in body size to ontogenetic growth and inheritance, where the latter captures the combined effects of plasticity, maternal effects and heritability. We show how these apparent discrepancies are mostly due to differences in aims and definitions. For example, the APE and IPM capture selection, whereas the AM and GM focus on the response to selection. Furthermore, the frameworks differ in the processes that are ascribed to plasticity and in how they take into account demography. We conclude that no single framework provides the ‘true’ contributions of evolution, plasticity and demography. Instead, different research questions require different frameworks. A thorough understanding of the different definitions of their components is necessary for selecting the most appropriate framework for the question at hand and for making biologically meaningful inferences. This work thus supports both future analysis and the careful interpretation of existing work.

Although the pedigree-based inbreeding coefficient F predicts the expected proportion of an individual's genome that is identical-by-descent (IBD), heterozygosity at genetic markers captures Mendelian sampling variation and thereby provides an estimate of realized IBD. Realized IBD should hence explain more variation in fitness than their pedigree-based expectations, but how many markers are required to achieve this in practice remains poorly understood. We use extensive pedigree and life-history data from an island population of song sparrows (Melospiza melodia) to show that the number of genetic markers and pedigree depth affected the explanatory power of heterozygosity and F, respectively, but that heterozygosity measured at 160 microsatellites did not explain more variation in fitness than F This is in contrast with other studies that found heterozygosity based on far fewer markers to explain more variation in fitness than F Thus, the relative performance of marker- and pedigree-based estimates of IBD depends on the quality of the pedigree, the number, variability and location of the markers employed, and the species-specific recombination landscape, and expectations based on detailed and deep pedigrees remain valuable until we can routinely afford genotyping hundreds of phenotyped wild individuals of genetic non-model species for thousands of genetic markers.

AbstractIn natural populations, quantitative trait dynamics often do not appear to follow evolutionary predictions: Despite abundant examples of natural selection acting on heritable traits, conclusive evidence for contemporary adaptive evolution remains rare for wild vertebrate populations, and phenotypic stasis seems to be the norm. This so-called ‘stasis paradox’ highlights our inability to predict evolutionary change, which is especially concerning within the context of rapid anthropogenic environmental change. While the causes underlying the stasis paradox are hotly debated, comprehensive attempts aiming at a resolution are lacking. Here we apply a quantitative genetic framework to individual-based long-term data for a wild rodent population and show that despite a positive association between body mass and fitness, there has been a genetic change towards lower body mass. The latter represents an adaptive response to viability selection favouring juveniles growing up to become relatively small adults, i.e. with a low potential adult mass, which presumably complete their development earlier. This selection is particularly strong towards the end of the snow-free season, and it has intensified in recent years, coinciding which a change in snowfall patterns. Importantly, neither the negative evolutionary change, nor the selective pressures that drive it, are apparent on the phenotypic level, where they are masked by phenotypic plasticity and a non-causal (i.e. non-genetic) positive association between body mass and fitness, respectively. Estimating selection at the genetic level thereby enabled us to uncover adaptive evolution in action, and to identify the corresponding phenotypic selective pressure. We thereby demonstrate that natural populations can show a rapid and adaptive evolutionary response to a novel selective pressure, and that explicitly (quantitative) genetic models are able to provide us with an understanding of the causes and consequences of selection that is superior to purely phenotypic estimates of selection and evolutionary change.

Empirical evidence for diminishing fitness returns of beneficial mutations supports Fisher’s geometric model. We show that a similar pattern emerges through the phenomenon of regression to the mean and that few studies correct for it. Although biases are often small, regression to the mean has overemphasized diminishing returns and will hamper cross-study comparisons unless corrected for.

Maternal hormones are important mediators of prenatal maternal effects. Although many experimental studies have demonstrated their potency in shaping offspring phenotypes, we know remarkably little about their adaptive value. Using long-term data on a wild collared flycatcher (Ficedula albicollis) population, we show that natural selection acts in opposite ways on two maternally derived androgens, yolk androstenedione (A4) and yolk testosterone (T). High yolk A4 concentrations are associated with higher fitness, whereas high yolk T concentrations are associated with lower fitness. Natural selection thus favours females that produce eggs with high A4 and low T concentrations. Importantly, however, there exists a positive (non-genetic) correlation between A4 and T, which suggests that females are limited in their ability to reach this adaptive optimum. Thereby, these results provide strong evidence for an adaptive value of differential maternal androgen deposition, and a mechanistic explanation for the maintenance of variation in maternal investment in the wild.

Egg components are important mediators of prenatal maternal effects in birds and other oviparous species. Because different egg components can have opposite effects on offspring phenotype, selection is expected to favour their mutual adjustment, resulting in a significant covariation between egg components within and/or among clutches. Here we tested for such correlations between maternally derived yolk immunoglobulins and yolk androgens in great tit (Parus major) eggs using a multivariate mixed-model approach. We found no association between yolk immunoglobulins and yolk androgens within clutches, indicating that within clutches the two egg components are deposited independently. Across clutches, however, there was a significant negative relationship between yolk immunoglobulins and yolk androgens, suggesting that selection has co-adjusted their deposition. Furthermore, an experimental manipulation of ectoparasite load affected patterns of covariance among egg components. Yolk immunoglobulins are known to play an important role in nestling immune defence shortly after hatching, whereas yolk androgens, although having growth-enhancing effects under many environmental conditions, can be immunosuppressive. We therefore speculate that variation in the risk of parasitism may play an important role in shaping optimal egg composition and may lead to the observed pattern of yolk immunoglobulin and yolk androgen deposition across clutches. More generally, our case study exemplifies how multivariate mixed-model methodology presents a flexible tool to not only quantify, but also test patterns of (co)variation across different organisational levels and environments, allowing for powerful hypothesis testing in ecophysiology. © 2013 Springer-Verlag Berlin Heidelberg.

Quality differences between offspring sired by the social and by an extra-pair partner are usually assumed to have a genetic basis, reflecting genetic benefits of female extra-pair mate choice. In the zebra finch (Taeniopygia guttata), we identified a colour ornament that is under sexual selection and appears to have a heritable basis. Hence, by engaging in extra-pair copulations with highly ornamented males, females could, in theory, obtain genes for increased offspring attractiveness. Indeed, sons sired by extra-pair partners had larger ornaments, seemingly supporting the genetic benefit hypothesis. Yet, when comparing ornament size of the social and extra-pair partners, there was no difference. Hence, the observed differences most likely had an environmental basis, mediated, for example, via differential maternal investment of resources into the eggs fertilized by extra-pair and social partners. Such maternal effects may (at least partly) be mediated by egg size, which we found to be associated with mean ornament expression in sons. Our results are consistent with the idea that maternal effects can shape sexual selection by altering the genotype-phenotype relationship for ornamentation. They also caution against automatically attributing greater offspring attractiveness or viability to an extra-pair mate's superior genetic quality, as without controlling for differential maternal investment we may significantly overestimate the role of genetic benefits in the evolution of extra-pair mating behaviour.

Because selection is often sex-dependent, alleles can have positive effects on fitness in one sex and negative effects in the other, resulting in intralocus sexual conflict. Evolutionary theory predicts that intralocus sexual conflict can drive the evolution of sex limitation, sex-linkage, and sex chromosome differentiation. However, evidence that sex-dependent selection results in sex-linkage is limited. Here, we formally partition the contribution of Y-linked and non-Y-linked quantitative genetic variation in coloration, tail, and body size of male guppies (Poecilia reticulata)-traits previously implicated as sexually antagonistic. We show that these traits are strongly genetically correlated, both on and off the Y chromosome, but that these correlations differ in sign and magnitude between both parts of the genome. As predicted, variation in attractiveness was found to be associated with the Y-linked, rather than with the non-Y-linked component of genetic variation in male ornamentation. These findings show how the evolution of Y-linkage may be able to resolve sexual conflict. More generally, they provide unique insight into how sex-specific selection has the potential to differentially shape the genetic architecture of fitness traits across different parts of the genome. © 2011 the Author(s). Evolution © 2011 the Society for the Study of Evolution.

1. Efforts to understand the links between evolutionary and ecological dynamics hinge on our ability to measure and understand how genes influence phenotypes, fitness and population dynamics. Quantitative genetics provides a range of theoretical and empirical tools with which to achieve this when the relatedness between individuals within a population is known. 2. A number of recent studies have used a type of mixed-effects model, known as the animal model, to estimate the genetic component of phenotypic variation using data collected in the field. Here, we provide a practical guide for ecologists interested in exploring the potential to apply this quantitative genetic method in their research. 3. We begin by outlining, in simple terms, key concepts in quantitative genetics and how an animal model estimates relevant quantitative genetic parameters, such as heritabilities or genetic correlations. 4. We then provide three detailed example tutorials, for implementation in a variety of software packages, for some basic applications of the animal model. We discuss several important statistical issues relating to best practice when fitting different kinds of mixed models. 5. We conclude by briefly summarizing more complex applications of the animal model, and by highlighting key pitfalls and dangers for the researcher wanting to begin using quantitative genetic tools to address ecological and evolutionary questions.

The ease with which Zebra Finches can be kept and bred in captivity makes them a suitable model for avian quantitative genetic studies. After a brief introduction to some quantitative genetic concepts, we here provide an up-to-date overview of quantitative genetic studies in Zebra Finches. We discuss what these studies can teach us about the evolutionary and behavioural ecology of Zebra Finches and song birds in general, and make suggestions for future research. Throughout this article we plead for a greater appreciation of the advantages offered by working on captive birds, but also discuss their limitations. Although quantitative genetic analyses in natural populations are becoming increasingly powerful, these studies lack the control possible in captivity. However, obtaining meaningful estimates of the type and strength of selection acting on phenotypic variation is more difficult in captivity. Hence, quantitative genetic studies in the wild and captivity each have their strengths and weaknesses and should be considered complementary rather than opposing. However, whereas quantitative genetic studies in the wild have boomed, the unique advantages offered by captive Zebra Finches have remained underexploited. Here we make a first attempt at changing this by highlighting what we believe may be fruitful lines for future research. © 2010 Royal Australasian Ornithologists Union.

Divergent selection pressures among populations can result not only in significant differentiation in morphology, physiology and behaviour, but also in how these traits are related to each other, thereby driving the processes of local adaptation and speciation. In the Australian zebra finch, we investigated whether domesticated stock, bred in captivity over tens of generations, differ in their response to a life-history manipulation, compared to birds taken directly from the wild. In a 'common aviary' experiment, we thereto experimentally manipulated the environmental conditions experienced by nestlings early in life by means of a brood size manipulation, and subsequently assessed its short- and long-term consequences on growth, ornamentation, immune function and reproduction. As expected, we found that early environmental conditions had a marked effect on both short- and long-term morphological and life-history traits in all birds. However, although there were pronounced differences between wild and domesticated birds with respect to the absolute expression of many of these traits, which are indicative of the different selection pressures wild and domesticated birds were exposed to in the recent past, manipulated rearing conditions affected morphology and ornamentation of wild and domesticated finches in a very similar way. This suggests that despite significant differentiation between wild and domesticated birds, selection has not altered the relationships among traits. Thus, life-history strategies and investment trade-offs may be relatively stable and not easily altered by selection. This is a reassuring finding in the light of the widespread use of domesticated birds in studies of life-history evolution and sexual selection, and suggests that adaptive explanations may be legitimate when referring to captive bird studies.

AbstractThe ability to predict individual breeding values in natural populations with known pedigrees has provided a powerful tool to separate phenotypic values into their genetic and environmental components in a nonexperimental setting. This has allowed sophisticated analyses of selection, as well as powerful tests of evolutionary change and differentiation. To date, there has, however, been no evaluation of the reliability or potential limitations of the approach. In this article, I address these gaps. In particular, I emphasize the differences between true and predicted breeding values (PBVs), which as yet have largely been ignored. These differences do, however, have important implications for the interpretation of, firstly, the relationship between PBVs and fitness, and secondly, patterns in PBVs over time. I subsequently present guidelines I believe to be essential in the formulation of the questions addressed in studies using PBVs, and I discuss possibilities for future research.

. Theoretical and laboratory research suggests that phenotypic plasticity can evolve under selection. However, evidence for its evolutionary potential from the wild is lacking. We present evidence from a Dutch population of great tits (
. Parus major
. ) for variation in individual plasticity in the timing of reproduction, and we show that this variation is heritable. Selection favoring highly plastic individuals has intensified over a 32-year period. This temporal trend is concurrent with climate change causing a mismatch between the breeding times of the birds and their caterpillar prey. Continued selection on plasticity can act to alleviate this mismatch.
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