Genomic basis of melanin-associated phenotypes suggests colour-specific environmental adaptations in tawny owls.

Feathers comprise a series of evolutionary innovations but also harbour colour, a key biological trait known to co-vary with life history or complex traits. Those relationships are particularly true in melanin-based pigmentation species due to known pleiotropic effects of the melanocortin pathway - originating from melanin-associated phenotypes. Here, we explore the molecular basis of melanin colouration and expected co-variation at the molecular level in the melanin-based, colour polymorphic system of the tawny owl (Strix aluco). An extensive body of literature has revealed that grey and brown tawny owl colour morphs differ in a series of life history and behavioural traits. Thus, it is plausible to expect co-variation also at molecular level between colour morphs. To investigate this possibility, we assembled the first draft genome of the species against which we mapped ddRADseq reads from 220 grey and 150 brown morphs - representing 10 years of pedigree data from a population in Southern Finland - and explored genome-wide associations with colour phenotype. Our results revealed putative molecular signatures of cold adaptation strongly associated with the grey phenotype, namely, a non-synonymous substitution in MCHR1, plus 2 substitutions in non-coding regions of FTCD and FAM135A whose genotype combinations obtained a predictive power of up to 100% (predicting grey colour). These suggest a molecular basis of cold environment adaptations predicted to be grey-morph specific. Our results potentially reveal part of the molecular machinery of melanin-associated phenotypes and provide novel insights towards understanding the functional genomics of colour polymorphism in melanin-based pigmented species.

Camouflage efficiency in a colour-polymorphic predator is dependent on environmental variation and snow presence in the wild.

Colour polymorphism can be maintained by colour morph-specific benefits across environmental conditions. Currently, the amount and the duration of snow cover during winter decrease especially in northern latitudes, which can alter the potential for camouflage of animals with light and dark morphs. Tawny owls, Strix aluco, are colour-polymorphic avian predators with dark (brown) and light (grey) colour morphs, where the grey morph is presumed to enjoy camouflage benefits under snowy conditions. We studied the camouflage potential of morphs in two tawny owls potential using passerines' probability to mob in the wild during winter with and without snow. For comparison with other seasons, we also repeated the experiment during spring and autumn. We found that grey tawny owls have a lower probability of being mobbed than the brown tawny owls only during snowy winters. The two colour morphs therefore experience differential benefits across snow conditions, which may help to maintain colour morphs in the population, although further warming of winter climate will reduce the potential for camouflage for grey tawny owls in northern latitudes.

Factors promoting hunting groups' sustainable harvest of moose in a co-management system.

Collaboration between and within management levels and involvement of local communities (co-management) increases sustainable management of natural resources. In Finland, moose (Alces alces) are harvested by hunting groups within a co-management system, providing meat and social benefits. We computed the 14-year change in moose harvest (2007-2020) for 4320 hunting groups. Moose harvest declined on average 1.1% per year, but with substantial variation in moose harvest changes between the hunting groups. We extracted information describing the collaboration between the hunting groups, their democratic status as well as leader dynamics, and the year of establishment. A hunting group's moose harvest was more stable (i.e. declined less) when the hunting group was (1) established a longer time ago; (2), had more changes in leadership over time, but did not depend on collaboration with other local hunting groups (in terms of jointly holding moose hunting licenses), whether the hunting group was a registered society (presumed to be more democratic than a non-registered one) or had consecutive leaders that shared a surname (presumed to be related). We conclude that encouraging resource users' early establishment in groups and groups' long-term persistence and promoting democratic leadership roles improves stable benefits from a natural resource in a co-management system.

Density-habitat relationships of white-tailed deer (Odocoileus virginianus) in Finland.

In heterogeneous landscapes, resource selection constitutes a crucial link between landscape and population-level processes such as density. We conducted a non-invasive genetic study of white-tailed deer in southern Finland in 2016 and 2017 using fecal DNA samples to understand factors influencing white-tailed deer density and space use in late summer prior to the hunting season. We estimated deer density as a function of landcover types using a spatial capture-recapture (SCR) model with individual identities established using microsatellite markers. The study revealed second-order habitat selection with highest deer densities in fields and mixed forest, and third-order habitat selection (detection probability) for transitional woodlands (clear-cuts) and closeness to fields. Including landscape heterogeneity improved model fit and increased inferred total density compared with models assuming a homogenous landscape. Our findings underline the importance of including habitat covariates when estimating density and exemplifies that resource selection can be studied using non-invasive methods.

Bringing to light nuclear-mitochondrial insertions in the genomes of nocturnal predatory birds.

Mito-nuclear insertions, or NUMTs, relate to genetic material of mitochondrial origin that have been transferred to the nuclear DNA molecule. The increasing amounts of genomic data currently being produced presents an opportunity to investigate this type of patterns in genome evolution of non-model organisms. Identifying NUMTs across a range of closely related taxa allows one to generalize patterns of insertion and maintenance in autosomes, which is ultimately relevant to the understanding of genome biology and evolution. Here we collected existing pairwise genome-mitogenome data of the order Strigiformes, a group that includes all the nocturnal bird predators. We identified NUMTs by applying percent similarity thresholds after blasting mitochondrial genomes against nuclear genome assemblies. We identified NUMTsin all genomes with numbers ranging from 4 in Bubo bubo to 24 in Ciccaba nigrolineata. Statistical analyses revealed NUMT size to negatively correlate with NUMT's sequence similarity to with original mtDNA region. Lastly, characterizing these nuclear insertions of mitochondrial origin in a comparative genomics framework produced variable phylogenetic patterns, suggesting in some cases that insertions might pre-date speciation events within Strigiformes.

Decomposing the spatial and temporal effects of climate on bird populations in northern European mountains.

The relationships between species abundance or occurrence versus spatial variation in climate are commonly used in species distribution models to forecast future distributions. Under "space-for-time substitution", the effects of climate variation on species are assumed to be equivalent in both space and time. Two unresolved issues of space-for-time substitution are the time period for species' responses and also the relative contributions of rapid- versus slow reactions in shaping spatial and temporal responses to climate change. To test the assumption of equivalence, we used a new approach of climate decomposition to separate variation in temperature and precipitation in Fennoscandia into spatial, temporal, and spatiotemporal components over a 23-year period (1996-2018). We compiled information on land cover, topography, and six components of climate for 1756 fixed route surveys, and we modeled annual counts of 39 bird species breeding in the mountains of Fennoscandia. Local abundance of breeding birds was associated with the spatial components of climate as expected, but the temporal and spatiotemporal climatic variation from the current and previous breeding seasons were also important. The directions of the effects of the three climate components differed within and among species, suggesting that species can respond both rapidly and slowly to climate variation and that the responses represent different ecological processes. Thus, the assumption of equivalent species' response to spatial and temporal variation in climate was seldom met in our study system. Consequently, for the majority of our species, space-for-time substitution may only be applicable once the slow species' responses to a changing climate have occurred, whereas forecasts for the near future need to accommodate the temporal components of climate variation. However, appropriate forecast horizons for space-for-time substitution are rarely considered and may be difficult to reliably identify. Accurately predicting change is challenging because multiple ecological processes affect species distributions at different temporal scales.

Structural equation modeling reveals decoupling of ecological and self-perceived outcomes in a garden box social-ecological system.

It is well known that green urban commons enhance mental and physical well-being and improve local biodiversity. We aim to investigate how these outcomes are related in an urban system and which variables are associated with better outcomes. We model the outcomes of an urban common-box gardening-by applying the Social-Ecological Systems (SES) framework. We expand the SES framework by analyzing it from the perspective of social evolution theory. The system was studied empirically through field inventories and questionnaires and modeled quantitatively by Structural Equation Modeling (SEM). This method offers powerful statistical models of complex social-ecological systems. Our results show that objectively evaluated ecological outcomes and self-perceived outcomes are decoupled: gardening groups that successfully govern the natural resource ecologically do not necessarily report many social, ecological, or individual benefits, and vice versa. Social capital, box location, gardener concerns, and starting year influenced the changes in the outcomes. In addition, the positive association of frequent interactions with higher self-perceived outcomes, and lack of such association with relatedness of group members suggests that reciprocity rather than kin selection explains cooperation. Our findings exemplify the importance of understanding natural resource systems at a very low "grassroot" level.

Cold winters have morph-specific effects on natal dispersal distance in a wild raptor.

Dispersal is a key process with crucial implications in spatial distribution, density, and genetic structure of species' populations. Dispersal strategies can vary according to both individual and environmental features, but putative phenotype-by-environment interactions have rarely been accounted for. Melanin-based color polymorphism is a phenotypic trait associated with specific behavioral and physiological profiles and is, therefore, a good candidate trait to study dispersal tactics in different environments. Here, using a 40 years dataset of a population of color polymorphic tawny owls (Strix aluco), we investigated natal dispersal distance of recruiting gray and pheomelanic reddish-brown (hereafter brown) color morphs in relation to post-fledging winter temperature and individual characteristics. Because morphs are differently sensitive to cold winters, we predicted that morphs' natal dispersal distances vary according to winter conditions. Winter temperature did not affect the proportion of brown (or gray) among recruits. We found that dispersal distances correlate with winter temperature in an opposite manner in the two morphs. Although the gray morph undertakes larger movements in harsher conditions, likely because it copes better with winter severity, the brown morph disperses shorter distances when winters are harsher. We discuss this morph-specific natal dispersal pattern in the context of competition for territories between morphs and in terms of costs and benefits of these alternative strategies. Our results stress the importance of considering the interaction between phenotype and environment to fully disentangle dispersal movement patterns and provide further evidence that climate affects the behavior and local distribution of this species.

Protected area characteristics that help waterbirds respond to climate warming.

Protected area networks help species respond to climate warming. However, the contribution of a site's environmental and conservation-relevant characteristics to these responses is not well understood. We investigated how composition of nonbreeding waterbird communities (97 species) in the European Union Natura 2000 (N2K) network (3018 sites) changed in response to increases in temperature over 25 years in 26 European countries. We measured community reshuffling based on abundance time series collected under the International Waterbird Census relative to N2K sites' conservation targets, funding, designation period, and management plan status. Waterbird community composition in sites explicitly designated to protect them and with management plans changed more quickly in response to climate warming than in other N2K sites. Temporal community changes were not affected by the designation period despite greater exposure to temperature increase inside late-designated N2K sites. Sites funded under the LIFE program had lower climate-driven community changes than sites that did not received LIFE funding. Our findings imply that efficient conservation policy that helps waterbird communities respond to climate warming is associated with sites specifically managed for waterbirds.

Las redes de áreas protegidas ayudan a las especies a responder al calentamiento climático. Sin embargo, se sabe muy poco sobre la contribución de las características ambientales y relevantes para la conservación de un sitio a estas respuestas. Investigamos cómo la composición de las comunidades no reproductivas de aves acuáticas (97 especies) en la red (3,018 sitios) Natura 2000 de la Unión Europea (N2K) cambió en respuesta a los incrementos de la temperatura durante más de 25 años en 26 países europeos. Medimos la reorganización comunitaria con base en series temporales de abundancia recolectadas durante el Censo Internacional de Aves Acuáticas en relación con los objetivos de conservación de los sitios N2K, el periodo de asignación de fondos y el estado del plan de manejo. La composición comunitaria de las aves acuáticas en los sitios con planes de manejo y designados explícitamente para su protección cambió más rápidamente en respuesta al calentamiento climático que en otros sitios N2K. Los cambios comunitarios temporales no se vieron afectados por el periodo de asignación a pesar de una mayor exposición al incremento de la temperatura dentro de los sitios N2K de asignación tardía. Los sitios financiados por el programa LIFE tuvieron menos cambios comunitarios causados por el clima que los sitios que no recibieron este financiamiento. Nuestros hallazgos sugieren que la política de conservación eficiente que ayuda a las comunidades de aves acuáticas a responder al calentamiento climático está asociada con sitios específicamente gestionados para las aves acuáticas.

Estimating preharvest density, adult sex ratio, and fecundity of white-tailed deer using noninvasive sampling techniques.

Adult sex ratio and fecundity (juveniles per female) are key population parameters in sustainable wildlife management, but inferring these requires abundance estimates of at least three age/sex classes of the population (male and female adults and juveniles). Prior to harvest, we used an array of 36 wildlife camera traps during 2 and 3 weeks in the early autumn of 2016 and 2017, respectively. We recorded white-tailed deer adult males, adult females, and fawns from the pictures. Simultaneously, we collected fecal DNA (fDNA) from 92 20 m × 20 m plots placed in 23 clusters of four plots between the camera traps. We identified individuals from fDNA samples with microsatellite markers and estimated the total sex ratio and population density using spatial capture-recapture (SCR). The fDNA-SCR analysis concluded equal sex ratio in the first year and female bias in the second year, and no difference in space use between sexes (fawns and adults combined). Camera information was analyzed in a spatial capture (SC) framework assuming an informative prior for animals' space use, either (a) as estimated by fDNA-SCR (same for all age/sex classes), (b) as assumed from the literature (space use of adult males larger than adult females and fawns), or (c) by inferring adult male space use from individually identified males from the camera pictures. These various SC approaches produced plausible inferences on fecundity, but also inferred total density to be lower than the estimate provided by fDNA-SCR in one of the study years. SC approaches where adult male and female were allowed to differ in their space use suggested the population had a female-biased adult sex ratio. In conclusion, SC approaches allowed estimating the preharvest population parameters of interest and provided conservative density estimates.

Regime shift tipping point in hare population collapse associated with climatic and agricultural change during the very early 20th century.

Animal populations at northern latitudes may have cyclical dynamics that are degraded by climate change leading to trophic cascade. Hare populations at more southerly latitudes are characterized by dramatic declines in abundance associated with agricultural intensification. We focus on the impact of historical climatic and agricultural change on a mid-latitude population of mountain hares, Lepus timidus hibernicus. Using game bag records from multiple sites throughout Ireland, the hare population index exhibited a distinct regime shift. Contrary to expectations, there was a dynamical structure typical of northern latitude hare populations from 1853 to 1908, during which numbers were stable but cyclic with a periodicity of 8 years. This regime was replaced by dynamics more typical of southern latitude hare populations from 1909 to 1970, in which cycles were lost and numbers declined dramatically. Destabilization of the autumn North Atlantic Oscillation (NAO) led to the collapse of similar cycles in the hare population, coincident with the onset of agricultural intensification (a shift from small-to-large farms) in the first half of the 20th century. Similar, but more recent regime shifts have been observed in Arctic ecosystems and attributed to anthropogenic climate change. The present study suggests such shifts may have occurred at lower latitudes more than a century ago during the very early 20th century. It seems likely that similar tipping points in the population collapse of other farmland species may have occurred similarly early but went undocumented. As northern systems are increasingly impacted by climate change and probable expansion of agriculture, the interaction of these processes is likely to disrupt the pulsed flow of resources from cyclic populations impacting ecosystem function.

Benefits of protected areas for nonbreeding waterbirds adjusting their distributions under climate warming.

Climate warming is driving changes in species distributions and community composition. Many species have a so-called climatic debt, that is, shifts in range lag behind shifts in temperature isoclines. Inside protected areas (PAs), community changes in response to climate warming can be facilitated by greater colonization rates by warm-dwelling species, but also mitigated by lowering extirpation rates of cold-dwelling species. An evaluation of the relative importance of colonization-extirpation processes is important to inform conservation strategies that aim for both climate debt reduction and species conservation. We assessed the colonization-extirpation dynamics involved in community changes in response to climate inside and outside PAs. To do so, we used 25 years of occurrence data of nonbreeding waterbirds in the western Palearctic (97 species, 7071 sites, 39 countries, 1993-2017). We used a community temperature index (CTI) framework based on species thermal affinities to investigate species turnover induced by temperature increase. We determined whether thermal community adjustment was associated with colonization by warm-dwelling species or extirpation of cold-dwelling species by modeling change in standard deviation of the CTI (CTISD ). Using linear mixed-effects models, we investigated whether communities in PAs had lower climatic debt and different patterns of community change than communities outside PAs. For CTI and CTISD combined, communities inside PAs had more species, higher colonization, lower extirpation, and lower climatic debt (16%) than communities outside PAs. Thus, our results suggest that PAs facilitate 2 independent processes that shape community dynamics and maintain biodiversity. The community adjustment was, however, not sufficiently fast to keep pace with the large temperature increases in the central and northeastern western Palearctic. Our results underline the potential of combining CTI and CTISD metrics to improve understanding of the colonization-extirpation patterns driven by climate warming.

Beneficios de las Áreas Protegidas para las Aves Acuáticas No Reproductoras que Están Ajustando su Distribución Debido al Calentamiento Climático Resumen El calentamiento climático está generando cambios en la distribución y en la composición comunitaria de las especies. Muchas de ellas tienen una deuda climática, es decir, los cambios en la distribución se atrasan con respecto a los cambios en las isoclinas térmicas. Dentro de las áreas protegidas (APs), los cambios comunitarios como respuesta al calentamiento climático pueden facilitarse mediante tasas mayores de colonización por especies de climas cálidos, pero también pueden mitigarse al reducir las tasas de extirpación de las especies de climas fríos. Se requiere una evaluación de la importancia relativa de los procesos de colonización-extirpación para orientar las estrategias de conservación que buscan la reducción de la deuda climática y la conservación de las especies. Analizamos las dinámicas de colonización-extirpación que participan en los cambios comunitarios como respuesta al clima dentro y fuera de las APs. Para realizar lo anterior, usamos datos tomados durante 25 años de la presencia de aves acuáticas no reproductoras en el Paleártico occidental (97 especies, 7,071 sitios, 39 países, 1993-2017). Usamos un marco de trabajo del índice de temperatura comunitaria (ITC) basado en las afinidades térmicas de las especies para así investigar la rotación de especies inducida por el incremento en la temperatura. Determinamos si el ajuste térmico en la comunidad estuvo asociado con la colonización por especies de climas cálidos o con la extirpación de especies de climas fríos al modelar el cambio mediante una desviación estándar del ITC (ITCDS ). Con los modelos lineales de efectos mixtos investigamos si las comunidades dentro de las APs tenían una deuda climática más baja y patrones diferentes de cambio comunitario que las comunidades localizadas fuera de las APs. Con la combinación del ITC y deL ITCDS , las comunidades dentro de las APs tuvieron más especies, una mayor colonización, una menor extirpación y una deuda climática más baja (16%) que las comunidades fuera de las APs. Por lo tanto, nuestros resultados sugieren que las APs facilitan dos procesos independientes que moldean las dinámicas comunitarias y mantienen la biodiversidad. Sin embargo, el ajuste comunitario no fue lo suficientemente rápido para mantener el paso de los grandes incrementos en la temperatura de las regiones central y noreste del Paleártico occidental. Nuestros resultados resaltan el potencial que tiene la combinación de las medidas del ITC y del ICTDS para mejorar el entendimiento de los patrones de colonización-extirpación causados por el calentamiento climático.

Lining the nest with more feathers increases offspring recruitment probability: Selection on an extended phenotype in the blue tit.

Birds, among various other taxa, construct nests. Nests form an extended phenotype of the individual building it. Nests are used to extend control over the conditions in which offspring develop, and are therefore commonly considered to be shaped by selection. Nevertheless, scarcely any scientific evidence exist that nest composition is under selection. Here, we demonstrate with data from over 400 blue tit (Cyanistes caeruleus) nests collected over 8 years that a higher proportion of feathers in the nest lining is positively associated with the probability of offspring to recruit as a breeding adult later in life. Strikingly, the extended phenotype (nest) was associated stronger with recruitment probability than phenotypic traits that have typically been considered important in selection (laying date, and female size and condition). Our findings suggest that the choice of nest material could be a maternal behavior with potential lifelong effects on her offspring.

Fledging Mass Is Color Morph Specific and Affects Local Recruitment in a Wild Bird.

AbstractEarly-life conditions may have long-lasting effects on life history. In color polymorphic species, morph-specific sensitivity to environmental conditions may lead to differential fitness. In tawny owls (Strix aluco), pheomelanin-based color polymorphism is expected to be maintained because the brown morph has higher adult fitness in warmer environments, while selection favors the gray morph under colder conditions. Here we investigate body mass at fledging and its consequences until adulthood in a population at the species' cold range margin. Using 40 years of data (1979-2017), we show that brown pairs, which mainly produce brown offspring consistent with a one-locus-two-alleles inheritance model, consistently raised heavier offspring than mixed (gray-brown) pairs and gray pairs. Offspring mass declined seasonally, except among offspring raised by brown pairs. Brown offspring could be heavier because of morph-specific parental care and/or offspring growth. Furthermore, mass at fledging is associated with fitness: the probability of local recruitment into the breeding population increased with higher mass at fledging, especially in mild winters and with favorable food conditions, although recruitment is not morph specific. Fledgling mass thus provides a fitness benefit in terms of recruitment probability that is modulated by environmental factors, which appear to level off any direct morph-specific recruitment benefits.

Bats and Wind Farms: The Role and Importance of the Baltic Sea Countries in the European Context of Power Transition and Biodiversity Conservation.

Although labeled as environmentally friendly, wind power can have negative impacts on the environment, such as habitat destruction or wildlife fatalities. Considering the distribution and migratory characteristics of European bats, the negative effects of wind power should be addressed on an appropriate scale. This review summarizes the current state of knowledge on interactions between wind farms and bats in Europe, and compares it with the situation in the countries of the European boreal biogeographic region. We analyzed data from papers published in international and national scientific journals, focusing on studies conducted in Europe. The issue of the impacts wind power has on bats is clearly overlooked in most of the countries of the European boreal region, with low volumes of research available on the topic. This is probably due to fewer wind farms in the area, making this recent issue a less-prioritized topic. However, the Baltic Sea, and the countries surrounding it, are of extreme importance with regards to bat migration, especially for the Pipistrellus nathusii. Therefore, more research on wind power and bats is needed in this region, as well as more cooperation between all the stakeholders.

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

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

Within-season changes in habitat use of forest-dwelling boreal bats.

Bats utilize forests as roosting sites and feeding areas. However, it has not been documented how bats utilize these habitats in the boreal zone with methods afforded by recent technological advances. Forest structure and management practices can create a variety of three-dimensional habitats for organisms capable of flight, such as bats. Here, we study the presence of boreal bats in a forest forming a mosaic of different age classes, dominant tree species, canopy cover, soil fertility, and other environmental variables, throughout their active season in the summer using passive ultrasound detectors. Our results indicate a preference for mature forest by Eptesicus nilssonii and a pooled set of Myotis bats. Both groups of bats also showed temporal changes in their habitat use regarding forest age. In June and July, both groups occurred more often in mature than young forests, but from August onwards, the difference in occurrence became less evident in Myotis and disappeared completely in E. nilssonii. In addition, E. nilssonii was more often present in forests with low canopy cover, and its occurrence shifted from coniferous forests to deciduous forests during the season. The results reflect the within-season dynamics of bat communities and their ability to utilize different types of forest as environmental conditions change. Yet, the results most importantly emphasize the importance of mature forests to bat diversity and the need to conserve such environments in the boreal zone.

Exploratory behavior undergoes genotype-age interactions in a wild bird.

Animal personality traits are often heritable and plastic at the same time. Indeed, behaviors that reflect an individual's personality can respond to environmental factors or change with age. To date, little is known regarding personality changes during a wild animals' lifetime and even less about stability in heritability of behavior across ages. In this study, we investigated age-related changes in the mean and in the additive genetic variance of exploratory behavior, a commonly used measure of animal personality, in a wild population of great tits. Heritability of exploration is reduced in adults compared to juveniles, with a low genetic correlation across these age classes. A random regression animal model confirmed the occurrence of genotype-age interactions (G×A) in exploration, causing a decrease in additive genetic variance before individuals become 1 year old, and a decline in cross-age genetic correlations between young and increasingly old individuals. Of the few studies investigating G×A in behaviors, this study provides rare evidence for this phenomenon in an extensively studied behavior. We indeed demonstrate that heritability and cross-age genetic correlations in this behavior are not stable over an individual's lifetime, which can affect its potential response to selection. Because G×A is likely to be common in behaviors and have consequences for our understanding of the evolution of animal personality, more attention should be turned to this phenomenon in the future work.

Tail colour signals performance in blue tit nestlings.

Indirect sexual selection arises when reproductive individuals choose their mates based on heritable ornaments that are genetically correlated to fitness. Evidence for genetic associations between ornamental colouration and fitness remains scarce. In this study, we investigate the quantitative genetic relationship between different aspects of tail structural colouration (brightness, hue and UV chroma) and performance (cell-mediated immunity, body mass and wing length) in blue tit (Cyanistes caeruleus) nestlings. In line with previous studies, we find low heritability for structural colouration and moderate heritability for performance measures. Multivariate animal models show positive genetic correlations between the three measures of performance, indicating quantitative genetic variation for overall performance, and tail brightness and UV chroma, two genetically independent colour measures, are genetically correlated with performance (positively and negatively, respectively). Our results suggest that mate choice based on independent aspects of tail colouration can have fitness payoffs in blue tits and provide support for the indirect benefits hypothesis. However, low heritability of tail structural colouration implies that indirect sexual selection on mate choice for this ornament will be a weak evolutionary force.