Evaluating indirect genetic effects of siblings using singletons

Estimating effects of parental and sibling genotypes (indirect genetic effects) can provide insight into how the family environment influences phenotypic variation. There is growing molecular genetic evidence for effects of parental phenotypes on their offspring (e.g. parental educational attainment), but the extent to which siblings affect each other is currently unclear. Here we used data from samples of unrelated individuals, without (singletons) and with biological full-siblings (non-singletons), to investigate and estimate sibling effects. Indirect genetic effects of siblings increase (or decrease) the covariance between genetic variation and a phenotype. It follows that differences in genetic association estimates between singletons and non-singletons could indicate indirect genetic effects of siblings if there is no heterogeneity in other sources of genetic association between singletons and non-singletons. We used UK Biobank data to estimate polygenic score (PGS) associations for height, BMI and educational attainment in self-reported singletons (N = 50,143) and non-singletons (N = 328,549). The educational attainment PGS association estimate was 12% larger (95% C.I. 3%, 21%) in the non-singleton sample than in the singleton sample, but the height and BMI PGS associations were consistent. Birth order data suggested that the difference in educational attainment PGS associations was driven by individuals with older siblings rather than firstborns. The relationship between number of siblings and educational attainment PGS associations was non-linear; PGS associations were 24% smaller in individuals with 6 or more siblings compared to the rest of the sample (95% C.I. 11%, 38%). We estimate that a 1 SD increase in sibling educational attainment PGS corresponds to a 0.025 year increase in the index individual’s years in schooling (95% C.I. 0.013, 0.036). Our results suggest that older siblings may influence the educational attainment of younger siblings, adding to the growing evidence that effects of the environment on phenotypic variation partially reflect social effects of germline genetic variation in relatives.     

New perspectives on the evolution of exaggerated traits

The scaling of body parts is central to the evolution of morphology and shape. Most traits scale proportionally with each other and body size such that larger adults are essentially magnified versions of smaller ones. This pattern is so ubiquitous that departures from it – disproportionate scaling between trait and body size – pique interest because it can generate dramatically exaggerated traits. These extreme morphologies are frequently hypothesized to result from sexual selection and their study has a long history, with several hypotheses seeking to explain their evolution. Despite this effort, surprisingly little progress has been made in demonstrating the forms of selection that produce different scaling patterns or in identifying the mechanisms that underlie the expression and evolution of scaling relationships. Here we review recent insights regarding the proximate mechanisms that regulate and integrate trait growth and that offer a new framework for studying the evolution of morphological scaling.     

Natural selection for earlier male arrival to breeding grounds through direct and indirect effects in a migratory songbird

For migratory birds, the earlier arrival of males to breeding grounds is often expected to have fitness benefits. However, the selection differential on male arrival time has rarely been decomposed into the direct effect of male arrival and potential indirect effects through female traits. We measured the directional selection differential on male arrival time in the pied flycatcher (Ficedula hypoleuca) using data from 6 years and annual number of fledglings as the fitness proxy. Using structural equation modeling, we were able to take into account the temporal structure of the breeding cycle and the hierarchy between the examined traits. We found directional selection differentials for earlier male arrival date and earlier female laying date, as well as strong selection differential for larger clutch size. These selection differentials were due to direct selection only as indirect selection for these traits was nonsignificant. When decomposing the direct selection for earlier male arrival into direct and indirect effects, we discovered that it was almost exclusively due to the direct effect of male arrival date on fitness and not due to its indirect effects via female traits. In other words, we showed for the first time that there is a direct effect of male arrival date on fitness while accounting for those effects that are mediated by effects of the social partner. Our study thus indicates that natural selection directly favored earlier male arrival in this flycatcher population.     

Longitudinal analyses of correlated response efficiencies of fillet traits in Nile tilapia

Recent studies with Nile tilapia have shown divergent results regarding the possibility of selecting on morphometric measurements to promote indirect genetic gains in fillet yield (FY). The use of indirect selection for fillet traits is important as these traits are only measurable after harvesting. Random regression models are a powerful tool in association studies to identify the best time point to measure and select animals. Random regression models can also be applied in a multiple trait approach to analyze indirect response to selection, which would avoid the need to sacrifice candidate fish. Therefore, the aim of this study was to investigate the genetic relationships between several body measurements, weight and fillet traits throughout the growth period and to evaluate the possibility of indirect selection for fillet traits in Nile tilapia. Data were collected from 2042 fish and was divided into two subsets. The first subset was used to estimate genetic parameters, including the permanent environmental effect for BW and body measurements (8758 records for each body measurement, as each fish was individually weighed and measured a maximum of six times). The second subset (2042 records for each trait) was used to estimate genetic correlations and heritabilities, which enabled the calculation of correlated response efficiencies between body measurements and the fillet traits. Heritability estimates across ages ranged from 0.05 to 0.5 for height, 0.02 to 0.48 for corrected length (CL), 0.05 to 0.68 for width, 0.08 to 0.57 for fillet weight (FW) and 0.12 to 0.42 for FY. All genetic correlation estimates between body measurements and FW were positive and strong (0.64 to 0.98). The estimates of genetic correlation between body measurements and FY were positive (except for CL at some ages), but weak to moderate (−0.08 to 0.68). These estimates resulted in strong and favorable correlated response efficiencies for FW and positive, but moderate for FY. These results indicate the possibility of achieving indirect genetic gains for FW and by selecting for morphometric traits, but low efficiency for FY when compared with direct selection.     

Darwinian balancing selection: Predation counters sexual selection in a wild insect

The potential viability costs of sexually selected traits are central to hypotheses about the evolution of exaggerated traits. Estimates of these costs in nature can come from selection analyses using multiple components of fitness during the same time frame. For a population of tree crickets (Oecanthus nigricornis: Gryllidae), we analyzed viability and sexual selection on male traits by comparing Oecanthus prey of a solitary wasp to those that survived, and comparing mating individuals to solitary males. We measured forewing width (sexually size dimorphic and used for singing), head width, pronotum length, and size of hind jumping legs as potential targets of selection. Supporting the hypothesis that sexually selected traits have viability costs, we found that significant directional sexual selection for wider heads was opposed by significant viability selection for narrower heads. Nonlinear selection revealed that individuals with wide heads and small legs were most attractive, but individuals with narrow heads, large legs, and intermediate pronotum length were most likely to survive. Successful mating may put males at greater risk of predation, especially if copulation per se is risky. Such balancing selection in tree crickets may have constrained the evolution of sexual dimorphism in head size—a condition seen in other gryllids and orthopterans.     

Positive genetic correlation between brain size and sexual traits in male guppies artificially selected for brain size

Brain size is an energetically costly trait to develop and maintain. Investments into other costly aspects of an organism's biology may therefore place important constraints on brain size evolution. Sexual traits are often costly and could therefore be traded off against neural investment. However, brain size may itself be under sexual selection through mate choice on cognitive ability. Here, we use guppy (Poecilia reticulata) lines selected for large and small brain size relative to body size to investigate the relationship between brain size, a large suite of male primary and secondary sexual traits, and body condition index. We found no evidence for trade‐offs between brain size and sexual traits. Instead, larger‐brained males had higher expression of several primary and precopulatory sexual traits – they had longer genitalia, were more colourful and developed longer tails than smaller‐brained males. Larger‐brained males were also in better body condition when housed in single‐sex groups. There was no difference in post‐copulatory sexual traits between males from the large‐ and small‐brained lines. Our data do not support the hypothesis that investment into sexual traits is an important limiting factor to brain size evolution, but instead suggest that brain size and several sexual traits are positively genetically correlated.     

Courtship success and multivariate analysis of sexual selection on morphometric traits inDrosophila buzzatii (Diptera: Drosophilidae)

Using wild-reared flies, we examined sexual selection on five phenotypic traits (thorax length, wing length, wing width, head width, and face width) inDrosophila buzzatii, by scoring copulatory status in nine mass mating cages. Only male face width was identified as a direct target of sexual selection in an analysis of selection gradient, while indirect selection was present on all other studied traits, as expected from their correlations with face width. In contrast to males, there was no indication of selection in females. Nor was there evidence of assortative mating. The suggested direct selection on face width seems to take place during licking behavior of the courtship and might be related to courtship feeding. This study suggests that courtship success gives rise to indirect selection on body size.     

Designing an early selection morphological linear traits index for dressage in the Pura Raza Español horse

Making a morphological pre-selection of Pura Raza Español horses (PRE) for dressage is a challenging task within its current breeding program. The aim of our research was to design an early genetic selection morphological linear traits index to improve dressage performance, using 26 morphological linear traits and six dressage traits (walk, trot, canter, submission, general impression – partial scores – and total score) as selection criteria. The data set included morphological linear traits of 10 127 PRE (4159 males and 5968 females) collected between 2008 and 2013 (one record per horse) and 19 095 dressage traits of 1545 PRE (1476 males and 69 females; 12.4 records of average) collected between 2004 and 2014. A univariate animal model was applied to predict the breeding values (PBV). A partial least squares regression analysis was used to select the most predictive morphological linear traits PBV on the dressage traits PBV. According to the Wold Criterion, the 13 morphological linear traits (width of head, head–neck junction, upper neck line, neck–body junction, width of chest, angle of shoulder, lateral angle of knee, frontal angle of knee, cannon bone perimeter, length of croup, angle of croup, ischium–stifle distance and lateral hock angle) most closely related to total score PBV, partial scores PBV and gait scores PBV (walk, trot and canter) were selected. A multivariate genetic analysis was performed among the 13 morphological linear traits selected and the six dressage traits to estimate the genetic parameters. After it, the selection index theory was used to compute the expected genetic response using different strategies. The expected genetic response of total score PBV (0.76), partial scores PBV (0.04) and gait scores PBV (0.03) as selection objectives using morphological linear traits PBV as criteria selection were positive, but lower than that obtained using dressage traits PBV (1.80, 0.16 and 0.14 for total score PBV, partial scores PBV and gait scores PBV) or dressage traits PBV and morphological linear traits PBV (2.97, 0.16 and 0.15 for total score PBV, partial scores PBV and gait scores PBV), as selection criteria. This suggests that it is possible to preselect the PRE without dressage traits PBV using as selection criteria the morphological linear traits PBV, but the expected genetic response will be lower.     

Additive and Non-additive Genetic Correlations. II. Application to Data Obtained from a 12 × 12 Diallel Cross

The proposed genetic correlation analysis, that involves the partitioning of the overall genetic correlation into an additive and a non-additive component, has been applied to data obtained from a diallel experiment involving 12 white modified opaque-2 maize inbred lines. The correlation analysis provided an insight into possible indirect selection strategies for the improvement of inferior kernel quality traits associated with the opaque-2 gene. Direct selection for high yield and low vitreousness rating would provide an efficient selection strategy for the development of high-yielding modified opaque-2 maize hybrids with desirable endosperm traits. It was concluded that it is not necessary to conduct the density, hardness and breakability determinations.     

Simultaneous selection on vegetative and reproductive phenology in a perennial herb

The timing of different life‐history events is often correlated, and selection might only rarely be exerted independently on the timing of a single event. In plants, phenotypic selection has often been shown to favor earlier flowering. However, little is known about to what extent this selection acts directly versus indirectly via vegetative phenology, and if selection on the two traits is correlational. We estimated direct, indirect, and correlational phenotypic selection on vegetative and reproductive phenology over 3 years for flowering individuals of the perennial herb Lathyrus vernus. Direct selection favored earlier flowering and shorter timespans between leaf‐out and flowering in all years. However, early flowering was associated with early leaf‐out, and the direction of selection on leaf‐out day varied among years. As a result, selection on leaf‐out weakened selection for early flowering in one of the study years. We found no evidence of correlational selection. Our results highlight the importance of including temporally correlated traits when exploring selection on the phenology of seasonal events.     

Genetic Variation Associated with Differential Educational Attainment in Adults Has Anticipated Associations with School Performance in Children

Genome-wide association study results have yielded evidence for the association of common genetic variants with crude measures of completed educational attainment in adults. Whilst informative, these results do not inform as to the mechanism of these effects or their presence at earlier ages and where educational performance is more routinely and more precisely assessed. Single nucleotide polymorphisms exhibiting genome-wide significant associations with adult educational attainment were combined to derive an unweighted allele score in 5,979 and 6,145 young participants from the Avon Longitudinal Study of Parents and Children with key stage 3 national curriculum test results (SATS results) available at age 13 to 14 years in English and mathematics respectively. Standardised (z-scored) results for English and mathematics showed an expected relationship with sex, with girls exhibiting an advantage over boys in English (0.433 SD (95%CI 0.395, 0.470), p<10⁻¹⁰) with more similar results (though in the opposite direction) in mathematics (0.042 SD (95%CI 0.004, 0.080), p = 0.030). Each additional adult educational attainment increasing allele was associated with 0.041 SD (95%CI 0.020, 0.063), p = 1.79×10⁻⁰⁴ and 0.028 SD (95%CI 0.007, 0.050), p = 0.01 increases in standardised SATS score for English and mathematics respectively. Educational attainment is a complex multifactorial behavioural trait which has not had heritable contributions to it fully characterised. We were able to apply the results from a large study of adult educational attainment to a study of child exam performance marking events in the process of learning rather than realised adult end product. Our results support evidence for common, small genetic contributions to educational attainment, but also emphasise the likely lifecourse nature of this genetic effect. Results here also, by an alternative route, suggest that existing methods for child examination are able to recognise early life variation likely to be related to ultimate educational attainment.     

Sexual dimorphism in the tusked frog, Adelotus brevis (Anura:Myobatrachidae): the roles of natural and sexual selection

At least two adaptive processes can lead to the evolution of sexual dimorphism: sexual selection (e.g. male-male combat) or natural selection (e.g. dietary divergence). We investigated the adaptive significance of a distinctive pattern of sexual dimorphism in a south-eastern Australian frog, Adelotus brevis. Male Adelotus grow larger than female conspecifics, have larger heads relative to body size, and have large paired projections (‘tusks’) in the lower jaw. All of these traits are rare among anurans. We quantified the degree of dimorphism in Adelotus, and gathered data on diets and mating systems of this species to evaluate the possible roles of sexual selection and dietary divergence in favoring die evolution of these sexually dimorphic traits. Analysis of prey items in alimentary tracts revealed significant sex differences in prey types. For example, females ate proportionally more arthropods and fewer molluscs than did males. However, this difference is likely to be a secondary consequence of habitat differences between the sexes (due in turn to their different reproductive roles) rather than a selective force for the evolution of sexual dimorphism. Calling males spend their time in moist habitats where pondsnails are abundant, whereas females are more often encountered in the drier arthropod-rich woodlands. A three-year behavioural ecology study on a field population revealed that reproductive males engage in agonistic interactions, with the sexually dimorphic tusks used to attack rivals. Larger body size contributed to male reproductive success. Small males were excluded from calling sites and, among the calling males, larger animals had higher reproductive success (numbers of matings) than did smaller individuals. Hence, the atypical pattern of sexual dimorphism in Adelotus brevis seems to have resulted from sexual selection for larger body size and tusk size in males, in the context of male-male agonistic behaviour, rather than natural selection for ecological divergence between the sexes.     

Brain size,head size and behaviour of a passerine bird

A recent increase in comparative studies of the ecological and evolutionary consequences of brain size in birds and primates in particular have suggested that cognitive abilities constitute a central link. Surprisingly, there are hardly any intraspecific studies investigating how individuals differing in brain size behave, how such individuals are distributed and how brain size is related to life history and fitness components. Brain mass of the barn swallow Hirundo rustica was strongly predicted by external head volume, explaining 99.5% of the variance, allowing for repeatable estimates of head volume as a reflection of brain size. Repeatability of head volume within and between years was high, suggesting that measurement errors were small. In a 2 years study of 501 individual adult barn swallows, I showed that head volume differed between sexes and age classes, with yearlings having smaller and more variable heads than older individuals, and females having smaller and more variable heads than males. Large head volume was not a consequence of large body size, which was a poor predictor of head volume. Birds with large heads arrived early from spring migration, independent of sex and age, indicating that migratory performance may have an important cognitive component. Head volume significantly predicted capture date and recapture probability, suggesting that head volume is related to learning ability, although morphological traits such as wing length, aspect ratio and wing area were unimportant predictors. Intensity of defence of offspring increased with head volume in females, but not in males. Barn swallows with large heads aggregated in large colonies, suggesting that individuals with large heads were more common in socially complex environments. These results suggest that brain size is currently under natural and sexual selection, and that micro‐evolutionary processes affecting brain size can be studied under field conditions.     

Revising traditional theory on the link between plant body size and fitness under competition: evidence from old‐field vegetation

The selection consequences of competition in plants have been traditionally interpreted based on a “size‐advantage” hypothesis – that is, under intense crowding/competition from neighbors, natural selection generally favors capacity for a relatively large plant body size. However, this conflicts with abundant data, showing that resident species body size distributions are usually strongly right‐skewed at virtually all scales within vegetation. Using surveys within sample plots and a neighbor‐removal experiment, we tested: (1) whether resident species that have a larger maximum potential body size (MAX) generally have more successful local individual recruitment, and thus greater local abundance/density (as predicted by the traditional size‐advantage hypothesis); and (2) whether there is a general between‐species trade‐off relationship between MAX and capacity to produce offspring when body size is severely suppressed by crowding/competition – that is, whether resident species with a larger MAX generally also need to reach a larger minimum reproductive threshold size (MIN) before they can reproduce at all. The results showed that MIN had a positive relationship with MAX across resident species, and local density – as well as local density of just reproductive individuals – was generally greater for species with smaller MIN (and hence smaller MAX). In addition, the cleared neighborhoods of larger target species (which had relatively large MIN) generally had – in the following growing season – a lower ratio of conspecific recruitment within these neighborhoods relative to recruitment of other (i.e., smaller) species (which had generally smaller MIN). These data are consistent with an alternative hypothesis based on a ‘reproductive‐economy‐advantage’ – that is, superior fitness under competition in plants generally requires not larger potential body size, but rather superior capacity to recruit offspring that are in turn capable of producing grand‐offspring – and hence transmitting genes to future generations – despite intense and persistent (cross‐generational) crowding/competition from near neighbors. Selection for the latter is expected to favor relatively small minimum reproductive threshold size and hence – as a tradeoff – relatively small (not large) potential body size.     

Selfing ability and dispersal are positively related,but not affected by range position: a multispecies study on southern African Asteraceae

Dispersal and breeding system traits are thought to affect colonization success. As species have attained their present distribution ranges through colonization, these traits may vary geographically. Although several theories predict associations between dispersal ability, selfing ability and the relative position of a population within its geographic range, there is little theoretical or empirical consensus on exactly how these three variables are related. We investigated relationships between dispersal ability, selfing ability and range position across 28 populations of 13 annual, wind‐dispersed Asteraceae species from the Namaqualand region of South Africa. Controlling for phylogeny, relative dispersal ability – assessed from vertical fall time of fruits – was positively related to an index of autofertility – determined from hand‐pollination experiments. These findings support the existence of two discrete syndromes: high selfing ability associated with good dispersal and obligate outcrossing associated with lower dispersal ability. This is consistent with the hypothesis that selection for colonization success drives the evolution of an association between these traits. However, no general effect of range position on dispersal or breeding system traits was evident. This suggests selection on both breeding system and dispersal traits acts consistently across distribution ranges.     

Mental Health Problems and Educational Attainment in Adolescence: 9-Year Follow-Up of the TRAILS Study

BackgroundThis study examines if mental health problems at age 11 and changes in mental health problems between age 11 and 16 predict educational attainment of adolescents at age 19, overall and stratified by gender.MethodsData from 1711 adolescents (76.8% from initial cohort) of the Tracking Adolescents'' Individual Lives Survey (TRAILS), a Dutch prospective cohort study with 9year follow-up, were used. Mental health problems (externalizing, internalizing and attention problems) were measured by the Youth Self Report and the Child Behavior Checklist at ages 11 and 16. Difference scores for mental health problems between age 11 and 16 were calculated. Educational attainment was assessed at age 19.ResultsExternalizing, internalizing and attention problems at age 11 were significantly associated with low educational attainment at age 19 (crude model). When adjusted for demographic variables and the other mental health problems, only the association for attention problems remained significant (odds ratio (OR), 95% confidence interval: 3.19, 2.11–4.83). Increasing externalizing problems between age 11 and 16 also predicted low educational attainment at age 19 (OR 3.12, 1.83–5.32). Among girls, increasing internalizing problems between age 11 and 16 predicted low educational attainment (OR 2.21, 1.25–3.94). For boys, no significant association was found for increasing internalizing problems and low educational attainment. For increasing attention problems between age 11 and 16 no significant association with low educational attainment was found.ConclusionsExternalizing, internalizing and attention problems at age 11 and an increase of these problems during adolescence predicted low educational attainment at age 19. Early treatment of these mental health problems may improve educational attainment, and reduce socioeconomic health differences in adulthood.     

Sexual size and shape dimorphism and allometric scaling patterns in head traits in the New Zealand common gecko Woodworthia maculatus

Sexual dimorphism in shape and size is widespread across animal taxa and arises when natural or sexual selection operates differently on the sexes. Male and female common geckos (Woodworthia maculatus; formerly Hoplodactylus maculatus) in New Zealand do not appear to experience different viability selection pressure, nor do males appear to be under intense pre-copulatory sexual selection. It was therefore predicted that this species would be sexually monomorphic with regard to body size and the size and shape of the head. In line with the prediction, there was no sexual difference in head width, depth, or length or in lateral head shape. However, contrary to prediction, males had a larger body and lateral head size than females. This study suggests that males, at least on Maud Island, NZ, might be under stronger pre-copulatory sexual selection than previously recognized and thus have evolved larger heads (i.e. lateral head size) for use in male combat for females. Allometric scaling patterns do not differ between the sexes and suggest that head width and depth are under directional selection whereas lateral head size is under stabilizing selection. Diet ecology – an agent of natural selection common to both sexes – is likely largely responsible for the observed patterns of head size and shape and the lack of sexual dimorphism in them.     

Life history trait divergence among populations of a non‐palatable species reveals strong non‐trophic indirect effects of an abundant herbivore

When large herbivores exert selection on their prey plant species, co‐occurring, non‐prey species may experience selection through non‐trophic indirect effects. Such selection is likely common where herbivores are overabundant. Yet, empirical studies of non‐trophic indirect effects as drivers of non‐prey trait evolution are lacking. Here we test for adaptive shifts in life history traits in an unpalatable species, Arisaema triphyllum, a common forest perennial that is unique because it exhibits size‐dependent sex switching. We collected A. triphyllum from six sites that experience a gradient in abiotic stress caused by deer browse pressure on prey plant species that generate indirect effects. We grew A. triphyllum from these sites in a common garden for five years to evaluate life history predictions linking strong indirect effects and abiotic stress to changes in life history traits: flowering onset size threshold, female flowering size threshold, relative growth rate (RGR), biomass allocation, and asexual reproduction. Despite observed differences among phenotypes in the field, expression of flowering onset size threshold, biomass allocation, and asexual reproduction did not differ among the six populations in the garden, indicating common plastic responses. In contrast, A. triphyllum collected from sites experiencing the two highest deer impacts exhibited smaller female flowering size thresholds and the highest RGR. Responses in these traits support the predictions of adaptive divergence in response to indirect effects. Our results reinforce the idea that non‐trophic indirect effects of large herbivores can elicit evolutionary responses in some traits of non‐prey species. In general, life history traits of unpalatable species may be cryptically adapting to stressful indirect effects where large herbivores are overabundant.     

The relationship between male head size and harem size in the sexually dimorphic mountain stone weta Hemideina maori

Abstract 1. Tree weta are a group of large, flightless orthopterans with pronounced sexual dimorphism. Males have enlarged heads that are used in fighting for possession of cavities in trees or under rocks where females shelter during the day.
2. The fieldwork reported here examined the relationship between male head size and mating success in Hemideina maori , an alpine tree weta that shelters under rock slabs that have broken off isolated outcrops or tors.
3. The relationship between male head size and harem size in H. maori is not as clear-cut as thought previously. First, overall body size is a better predictor of male mating success than head size per se . Second, both body size and head size explained a relatively low percentage (19.8%) of the overall variation in mating success. Third, despite the intensity of directional selection being estimated to move the frequency distribution of head size and femur size 0.49 and 0.54 standard deviations from the mean, male heads and femurs were ≈ 2 mm smaller at the main study site than at a second site 100 m higher in elevation. A similar pattern was found for adult females. Additional surveys have indicated that body size in H. maori decreases with decreasing altitude, which is correlated with increasing night-time temperature.
4. Although there are reasons why natural selection might favour weta maturing earlier and at smaller body sizes in warmer environments, relatively large males would still have a mating advantage over smaller males under such conditions. This sexually dimorphic alpine insect might be a good example of the trade-offs and conflicting demands that sexual selection versus natural selection can place on organisms.     

Natural and sexual selection on a plumage signal of status and on morphology in house sparrows,Passer domesticus

Temporal patterns of natural and sexual selection on male badge size and body traits were studied in a population of house sparrows, Passer domesticus. Badge size was a heritable trait as revealed by a significant father-son regression. Survival during autumn dispersal and winter was not related to badge size or body traits in yearling male house sparrows. Badges that signal dominance status were affected positively by directional selection for mating. Adult male house sparrows suffered an opposing selection pressure on badge size during autumn. Contrary to males, female house sparrows did not experience significant directional or stabilizing selection on any body trait. Directional sexual selection on male badge size due to female choice moves male sparrows away from their survival optimum. Opposing directional natural selection on badge size due to autumn mortality caused by predation maintains a stable badge size.