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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.

Changing environmental conditions cause changes in the distributions of phenotypic traits in natural populations. However, determining the mechanisms responsible for these changes-and, in particular, the relative contributions of phenotypic plasticity versus evolutionary responses-is difficult. To our knowledge, no study has yet reported evidence that evolutionary change underlies the most widely reported phenotypic response to climate change: the advancement of breeding times. In a wild population of red deer, average parturition date has advanced by nearly 2 weeks in 4 decades. Here, we quantify the contribution of plastic, demographic, and genetic components to this change. In particular, we quantify the role of direct phenotypic plasticity in response to increasing temperatures and the role of changes in the population structure. Importantly, we show that adaptive evolution likely played a role in the shift towards earlier parturition dates. The observed rate of evolution was consistent with a response to selection and was less likely to be due to genetic drift. Our study provides a rare example of observed rates of genetic change being consistent with theoretical predictions, although the consistency would not have been detected with a solely phenotypic analysis. It also provides, to our knowledge, the first evidence of both evolution and phenotypic plasticity contributing to advances in phenology in a changing climate.

Quantitative genetic analyses require extensive measurements of phenotypic traits, a task that is often not trivial, especially in wild populations. On top of instrumental measurement error, some traits may undergo transient (i.e., nonpersistent) fluctuations that are biologically irrelevant for selection processes. These two sources of variability, which we denote here as measurement error in a broad sense, are possible causes for bias in the estimation of quantitative genetic parameters. We illustrate how in a continuous trait transient effects with a classical measurement error structure may bias estimates of heritability, selection gradients, and the predicted response to selection. We propose strategies to obtain unbiased estimates with the help of repeated measurements taken at an appropriate temporal scale. However, the fact that in quantitative genetic analyses repeated measurements are also used to isolate permanent environmental instead of transient effects requires that the information content of repeated measurements is carefully assessed. To this end, we propose to distinguish “short-term” from “long-term” repeats, where the former capture transient variability and the latter help isolate permanent effects. We show how the inclusion of the corresponding variance components in quantitative genetic models yields unbiased estimates of all quantities of interest, and we illustrate the application of the method to data from a Swiss snow vole population.

Evolutionary ecologists dating back to Darwin (1871) have sought to understand why males are larger than females in some species, and why females are the larger sex in others. Although the former is widespread in mammals, rodents and other small mammals usually exhibit low levels of sexual size dimorphism (SSD). Here, we investigate patterns of sexual dimorphism in 34 vole species belonging to the subfamily Arvicolinae in a phylogenetic comparative framework. We address the potential role of sexual selection and fecundity selection in creating sex differences in body size. No support was found for hyperallometric scaling of male body size to female body size. We observed a marginally significant relationship between SSD and the ratio of male to female home range size, with the latter being positively related to the level of intrasexual competition for mates. This suggests that sexual selection favours larger males. Interestingly, we also found that habitat type, but not mating system, constitutes a strong predictor of SSD. Species inhabiting open habitats – where males have extensive home ranges in order to gain access to as many females as possible – exhibit a higher mean dimorphism than species inhabiting closed habitats, where females show strong territoriality and an uniform distribution preventing males to adopt a territorial strategy for gaining copulations. Nonetheless, variation in the strength of sexual selection is not the only selective force shaping SSD in voles; we also found a positive association between female size and litter size across lineages. Assuming this relationship also exists within lineages (i.e. fecundity selection on female size), this suggests an additional role for variation in the strength of fecundity selection shaping interspecific differences in female size, and indirectly in SSD. Therefore our results suggest that different selective processes act on the sizes of males and females, but because larger size is favoured in both sexes, SSD is on average relatively small.

1. Demographic and evolutionary modelling approaches are critical to understanding and projecting species responses to global environmental changes. Population matrix models have been a favoured tool in demography, but until recently, they failed to account for short-term evolutionary changes. Evolutionary-explicit demographic models remain computationally intensive, difficult to use and have yet to be widely adopted for empirical studies. Researchers focusing on short-term evolution often favour individual-based simulations, which are more flexible but less transferable and computationally efficient. Limited communication between fields has led to differing perspectives on key issues, such as how life-history traits affect adaptation to environmental change. We develop a new EvoDemo hyperstate matrix population model (EvoDemo-Hyper MPM) that incorporates the genetic inheritance of quantitative traits, enabling fast computation of evolutionary and demographic dynamics. We evaluate EvoDemo-Hyper MPM against individual-based simulations and provide analytical approximations for adaptation rates across six distinct scales in response to selection. We show that different methods yield equivalent results for the same biological scenario, although semantic differences between fields may obscure these similarities. Our results demonstrate thatEvoDemo-Hyper MPM provides accurate, computationally efficient solutions, closely matching outcomes from individual-based simulations and analytical approximations under similar biological conditions. Adaptation rates per generation remain constant across species when selection acts on fertility but vary with other vital rates. Adaptation per time decreases with generation time unless selection targets adult survival, where intermediate life histories adapt fastest. Rates per generation, defined as the relative change in individual fitness, remain constant across species and vital rates.4. We discuss that no general prediction emerges about which species or life-history traits yield higher adaptation rates, as outcomes depend on life cycles, vital rates | 1645 VAN de WALLE et al.

Background The shape of the semicircular canals of the inner ear of living squamate reptiles has been used to infer phylogenetic relationships, body size, and life habits. Often these inferences are made without controlling for the effects of the other ones. Here we examine the semicircular canals of 94 species of extant limbed lepidosaurs using three-dimensional landmark-based geometric morphometrics, and analyze them in phylogenetic context to evaluate the relative contributions of life habit, size, and phylogeny on canal shape. Results Life habit is not a strong predictor of semicircular canal shape across this broad sample. Instead, phylogeny plays a major role in predicting shape, with strong phylogenetic signal in shape as well as size. Allometry has a limited role in canal shape, but inner ear size and body mass are strongly correlated. Conclusions Our wide sampling across limbed squamates suggests that semicircular canal shape and size are predominantly a factor of phylogenetic relatedness. Given the small proportion of variance in semicircular canal shape explained by life habit, it is unlikely that unknown life habit could be deduced from semicircular canal shape alone. Overall, semicircular canal size is a good estimator of body length and even better for body mass in limbed squamates. Semiaquatic taxa tend to be larger and heavier than non-aquatic taxa, but once body size and phylogeny are accounted for, they are hard to distinguish from their non-aquatic relatives based on bony labyrinth shape and morphology.

Heterogeneity in fitness components consists of fixed heterogeneity due to latent differences fixed throughout life (e.g., genetic variation) and dynamic heterogeneity generated by stochastic variation. Their relative magnitude is crucial for evolutionary processes, as only the former may allow for adaptation. However, the importance of fixed heterogeneity in small populations has recently been questioned. Using neutral simulations (NS), several studies failed to detect fixed heterogeneity, thus challenging previous results from mixed models (MM). To understand the causes of this discrepancy, we estimate the statistical power and false positive rate of both methods and apply them to empirical data from a wild rodent population. While MM show high false-positive rates if confounding factors are not accounted for, they have high statistical power to detect real fixed heterogeneity. In contrast, NS are also subject to high false-positive rates but always have low power. Indeed, MM analyses of the rodent population data show significant fixed heterogeneity in reproductive success, whereas NS analyses do not. We suggest that fixed heterogeneity may be more common than is suggested by NS and that NS are useful only if more powerful methods are not applicable and if they are complemented by a power analysis.

Phenotypic trait data play a central role in ecology and evolutionary research. The quality of trait data, and the findings of subsequent analyses, depend on the quality of measurement. However, most studies overlook measurement accuracy in their study designs. We investigated the repeatability of five frequently used linear measurements of avian traits: wing length, tarsus length, bill length, bill depth and bill width and the validity of proxies for three traits: bill surface area, structural body size and tarsus size, using species from the infra-order Meliphagides (honeyeaters, fairy wrens and their allies). Repeatability varied between traits and across species for a given trait: traits larger than 13 mm showed high repeatability compared with smaller traits. By incorporating microCT technology, we showed that the formula for the surface area of a cone, a widely used proxy of bill surface area, accurately describes bill surface area within species. Surface measurement of tarsus and wing lengths were valid proxies for underlying osteology. We recommend preliminary estimation of repeatability should be undertaken for individual traits prior to data collection, in order to design suitable protocols that improve data quality, while optimizing costs involved, particularly for traits < 13 mm.

Many studies ask whether young or older males are better at acquiring mates. Even so, how age affects reproductive success is still poorly understood because male age and mating history are confounded in most studies: older males usually have more mating experience. To what extent does mating history rather than age explain variation in male mating success? And how do mating history and male age determine paternity when there is also postcopulatory sexual selection? Here, we experimentally manipulated the mating history of old and young males in the eastern mosquitofish (Gambusia holbrooki). We then recorded male mating behavior and share of paternity (1259 offspring from 232 potential sires) when they competed for mates and fertilizations. Old males, and males with no mating experience, spent significantly more time approaching females, and attempting to mate, than did young males and those with greater mating experience. Male age and mating history interacted to affect paternity: old males benefited from having previous mating experience, but young males did not. Our results highlight that the age-related changes in male reproductive traits and in paternity that have been described in many taxa may be partly attributable to male mating history and not simply to age itself.

Hybridization is increasingly recognized as a significant evolutionary process, in particular because it can lead to introgression of genes from one species to another. A striking pattern of discordance in the amount of introgression between mitochondrial and nuclear markers exists such that substantial mitochondrial introgression is often found in combination with no or little nuclear introgression. Multiple mechanisms have been proposed to explain this discordance, including positive selection for introgressing mitochondrial variants, several types of sex-biases, drift, negative selection against introgression in the nuclear genome, and spatial expansion. Most of these hypotheses are verbal, and have not been quantitatively evaluated so far. We use individual-based, multilocus, computer simulations of secondary contact under a wide range of demographic and genetic scenarios to evaluate the ability of the different mechanisms to produce discordant introgression. Sex-biases and spatial expansions fail to produce substantial mito-nuclear discordance. Drift and nuclear selection can produce strong discordance, but only under a limited range of conditions. In contrast, selection on the mitochondrial genome produces strong discordance, particularly when dispersal rates are low. However, commonly used statistical tests have little power to detect this selection. Altogether, these results dismiss several popular hypotheses, and provide support for adaptive mitochondrial introgression.