Pollinator‐mediated selection in a specialized pollination system: matches and mismatches across populations

Most studies on pollinator‐mediated selection have been performed in generalized rather than specialized pollination systems. This situation has impeded evaluation of the extent to which selection acts on attraction or specialized key floral traits involved in the plant‐pollinator phenotypic interphase. We studied pollinator‐mediated selection in four populations of Nierembergia linariifolia, a self‐incompatible and oil‐secreting plant pollinated exclusively by oil‐collecting bees. We evaluated whether floral traits experience variable selection among populations and whether attraction and fit traits are heterogeneously selected across populations. Populations differed in every flower trait and selection was consistently observed for corolla size and flower shape, two traits involved in the first steps of the pollination process. However, we found no selection acting on mechanical‐fit traits. The observation that selection occurred upon attraction rather than mechanical‐fit traits, suggests that plants are not currently evolving fine‐tuned morphological adaptations to local pollinators and that phenotypic matching is not necessarily an expected outcome in this specialized pollination system.     

Pollinator‐mediated selection in a specialized hummingbird–Heliconia system in the Eastern Caribbean

Phenotypic matches between plants and their pollinators often are interpreted as examples of reciprocal selection and adaptation. For the two co‐occurring plant species, Heliconia bihai and H. caribaea in the Eastern Caribbean, we evaluated for five populations over 2 years the strength and direction of natural selection on corolla length and number of bracts per inflorescence. These plant traits correspond closely to the bill lengths and body masses of their primary pollinators, female or male purple‐throated carib hummingbirds (Eulampis jugularis). In H. bihai, directional selection for longer corollas was always significant with the exception of one population in 1 year, whereas selection on bract numbers was rare and found only in one population in 1 year. In contrast, significant directional selection for more bracts per inflorescence occurred in all three populations of the yellow morph and in two populations of the red morph of H. caribaea, whereas significant directional selection on corolla length occurred in only one population of the red morph and one population of the yellow morph. Selection for longer corollas in H. bihai may result from better mechanical fit, and hence pollination, by the long bills of female E. jugularis, their sole pollinator. In contrast, competition between males of E. jugularis for territories may drive selection for more bracts in H. caribaea. Competitive exclusion of female E. jugularis by territorial males also implicates pollinator competition as a possible ecological mechanism for trait diversification in these plants.     

Morph‐specific selection on floral traits in a polymorphic plant

Correlations between phenotypic traits are common in many organisms, but the relative importance of nonadaptive mechanisms and selection for the evolution and maintenance of such correlations are poorly understood. In polymorphic species, morphs may evolve quantitative differences in additional characters as a result of morph‐specific selection. The perennial rosette herb Primula farinosa is polymorphic for scape length. The short‐scaped morph is less damaged by grazers and seed predators but is more strongly pollen limited than the long‐scaped morph. We examined whether morph‐specific differences in biotic interactions are associated with differences in selection on two other traits affecting floral display (number of flowers and petal size) and on one trait likely to affect pollination efficiency (corolla tube width) in three P. farinosa populations. Differences in selection between morphs were detected in one population. In this population, selection for more flowers and larger petals was stronger in the short‐scaped than in the long‐scaped morph, and although there was selection for narrower corolla tubes in the short‐scaped morph, no statistically significant selection on corolla tube width could be detected in the long‐scaped morph. In the study populations, the short‐scaped morph produced more and larger flowers and wider corolla tubes. Current morph‐specific selection was thus only partly consistent with trait differences between morphs. The results provide evidence of morph‐specific selection on traits associated with floral display and pollination efficiency, respectively.     

The origin of ecological divergence in Helianthus paradoxus (Asteraceae): selection on transgressive characters in a novel hybrid habitat

Diploid hybrid speciation in plants is often accompanied by rapid ecological divergence between incipient neospecies and their parental taxa. One plausible means by which novel adaptation in hybrid lineages may arise is transgressive segregation, that is, the generation of extreme phenotypes that exceed those of the parental lines. Early generation (BC2) hybrids between two wild, annual sunflowers, Helianthus annuus and Helianthus petiolaris, were used to study directional selection on transgressive characters associated with the origin of Helianthus paradoxus, a diploid hybrid species adapted to extremely saline marshes. The BC2 plants descended from a single F1 hybrid backcrossed toward H. petiolaris. The strength of selection on candidate adaptive traits in the interspecific BC2 was measured in natural H. paradoxus salt marsh habitat. Positive directional selection was detected for leaf succulence and Ca uptake, two traits that are known to be important in salt stress response in plants. Strong negative directional selection operated on uptake of Na and correlated elements. A significant decrease in trait correlations over time was observed in the BC2 population for Na and Ca content, suggesting an adaptive role for increased Ca uptake coupled with increased net exclusion of Na from leaves. Patterns of directional selection in BC2 hybrids were concordant with character expression in the natural hybrid species, H. paradoxus, transplanted into the wild. Moreover, the necessary variation for generating the H. paradoxus phenotype existed only in the BC2 population, but not in samples of the two parental species, H. annuus and H. petiolaris. These results are consistent with the hypothesis that transgressive segregation of elemental uptake and leaf succulence contributed to the origin of salt adaptation in the diploid hybrid species H. paradoxus.     

Quantifying hummingbird preference for floral trait combinations: The role of selection on trait interactions in the evolution of pollination syndromes

Darwin recognized the flower's importance for the study of adaptation and emphasized that the flower's functionality reflects the coordinated action of multiple traits. Here we use a multitrait manipulative approach to quantify the potential role of selection acting on floral trait combinations underlying the divergence and maintenance of three related North American species of Silene (Caryophyllaceae). We artificially generated 48 plant phenotypes corresponding to all combinations of key attractive traits differing among the three Silene species (color, height, inflorescence architecture, flower orientation, and corolla‐tube width). We quantified main and interaction effects of trait manipulation on hummingbird visitation preference using experimental arrays. The main effects of floral display height and floral orientation strongly influenced hummingbird visitation, with hummingbirds preferring flowers held high above the ground and vertically to the sky. Hummingbirds also prefer traits in a nonadditive manner as multiple two‐way and higher order interaction effects were important predictors of hummingbird visitation. Contemporary trait combinations found in hummingbird pollinated S. virginica are mostly preferred. Our study demonstrates the likelihood of pollination syndromes evolving due to selection on trait combinations and highlights the importance of trait interactions in understanding the evolution of complex adaptations.     

Strong pollinator‐mediated selection for increased flower brightness and contrast in a deceptive orchid

Contrasting flower color patterns that putatively attract or direct pollinators toward a reward are common among angiosperms. In the deceptive orchid Anacamptis morio, the lower petal, which makes up most of the floral display, has a light central patch with dark markings. Within populations, there is pronounced variation in petal brightness, patch size, amount of dark markings, and contrast between patch and petal margin. We tested whether pollinators mediate selection on these color traits and on morphology (plant height, number of flowers, corolla size, spur length), and whether selection is consistent with facilitated or negative frequency‐dependent pollination. Pollinators mediated strong selection for increased petal brightness (Δβ_poll = 0.42) and contrast (Δβ_poll = 0.51). Pollinators also tended to mediate stabilizing selection on brightness (Δγ_poll = –0.27, n.s.) favoring the most common phenotype in the population. Selection for reduced petal brightness among hand‐pollinated plants indicated a fitness cost associated with brightness. The results demonstrate that flower color traits influence pollination success and seed production in A. morio, indicating that they affect attractiveness to pollinators, efficiency of pollen transfer, or both. The documented selection is consistent with facilitated pollination and selection for color convergence toward cooccurring rewarding species.     

The effect of polyploidy and hybridization on the evolution of floral colour in Nicotiana (Solanaceae)

Background and Aims Speciation in angiosperms can be accompanied by changes in floral colour that may influence pollinator preference and reproductive isolation. This study investigates whether changes in floral colour can accompany polyploid and homoploid hybridization, important processes in angiosperm evolution.Methods Spectral reflectance of corolla tissue was examined for 60 Nicotiana (Solanaceae) accessions (41 taxa) based on spectral shape (corresponding to pigmentation) as well as bee and hummingbird colour perception in order to assess patterns of floral colour evolution. Polyploid and homoploid hybrid spectra were compared with those of their progenitors to evaluate whether hybridization has resulted in floral colour shifts.Key Results Floral colour categories in Nicotiana seem to have arisen multiple times independently during the evolution of the genus. Most younger polyploids displayed an unexpected floral colour, considering those of their progenitors, in the colour perception of at least one pollinator type, whereas older polyploids tended to resemble one or both of their progenitors.Conclusions Floral colour evolution in Nicotiana is weakly constrained by phylogeny, and colour shifts do occur in association with both polyploid and homoploid hybrid divergence. Transgressive floral colour in N. tabacum has arisen by inheritance of anthocyanin pigmentation from its paternal progenitor while having a plastid phenotype like its maternal progenitor. Potentially, floral colour evolution has been driven by, or resulted in, pollinator shifts. However, those polyploids that are not sympatric (on a regional scale) with their progenitor lineages are typically not divergent in floral colour from them, perhaps because of a lack of competition for pollinators.     

Floral scent evolution in the section Pseudophrys: pollinator‐mediated selection or phylogenetic constraints?

• Sexually deceptive orchid species from the Mediterranean genus Ophrys usually interact with one or a few pollinator species by means of specific floral scents. In this study, we investigated the respective role of pollinator‐mediated selection and phylogenetic constraints in the evolution of floral scents in the section Pseudophrys.
• We built a phylogenetic tree of 19 Pseudophrys species based on three nuclear loci; we gathered a dataset on their pollination interactions from the literature and from our own field data; and we extracted and analysed their floral scents using solid phase microextraction and gas chromatography‐mass spectrometry. We then quantified the phylogenetic signal carried by floral scents and investigated the link between plant–pollinator interactions and floral scent composition using phylogenetic comparative methods.
• We confirmed the monophyly of the section Pseudophrys and demonstrated the existence of three main clades within this section. We found that floral scent composition is affected by both phylogenetic relationships among Ophrys species and pollination interactions, with some compounds (especially fatty acid esters) carrying a significant phylogenetic signal and some (especially alkenes and alkadienes) generating dissimilarities between closely related Pseudophrys pollinated by different insects.
• Our results show that in the section Pseudophrys, floral scents are shaped both by pollinator‐mediated selection and by phylogenetic constraints, but that the relative importance of these two evolutionary forces differ among compound classes, probably reflecting distinct selective pressures imposed upon behaviourally active and non‐active compounds.

     

Enantiostyly in Chamaecrista ramosa (Fabaceae‐Caesalpinioideae): floral morphology,pollen transfer dynamics and breeding system

Enantiostyly is a form of reciprocal herkogamy, in which floral morphs present reciprocal differences in the position of sexual elements, and occurs in monomorphic and dimorphic forms. This polymorphism maximises cross‐pollination and reduces self‐pollination, being very common within the subtribe Cassiinae (Fabaceae). Nevertheless, few studies have investigated the functionality of enantiostyly, particularly in this plant group. The present study aimed to investigate enantiostyly and its functionality in Chamaecrista ramosa, a monomorphic enantiostylous shrub, in an area of coastal vegetation in northeast Brazil. Pollen deposition and capture on the body of floral visitors, the relationship of these data with floral biology and breeding system, and morph ratio were evaluated. Pollen deposition and capture occurred in specific sites of the floral visitor body, showing the functionality of enantiostyly. The floral architecture, associated with the floral visitor behaviour, resulted in indirect pollen deposition on the floral visitor body. This occurred through a loop made by the pollen upon the inner petal surface, similar that generally reported for other Cassiinae. Chamaecrista ramosa is self‐compatible, although no fruit set was observed through spontaneous self‐pollination. The occurrence and number of floral morphs was similar within clumps. Enantiostyly seems to be advantageous for this species, as it results in efficient pollen capture and deposition, reduces the chances of autogamy and maximises intermorph pollen flow.     

Natural selection on floral volatile production in Penstemon digitalis: Highlighting the role of linalool

Natural selection is thought to have shaped the evolution of floral scent; however, unlike other floral characters, we have a rudimentary knowledge of how phenotypic selection acts on scent. We found that floral scent was under stronger selection than corolla traits such as flower size and flower color in weakly scented Penstemon digitalis. Our results suggest that to understand evolution in floral phenotypes, including scent in floral selection, studies are crucial. For P. digitalis, linalool was the direct target of selection in the scent bouquet. Therefore, we determined the enantiomeric configuration of linalool because interacting insects may perceive the enantiomers differentially. We found that P. digitalis produces only (S)-(+)-linalool and, more interestingly, it is also taken up into the nectar. Because the nectar is scented and flavored with (S)-(+)-linalool, it may be an important cue for pollinators visiting P. digitalis flowers.     

Deconstructing a floral phenotype: do pollinators select for corolla integration in Lavandula latifolia?

An experimental field study was undertaken to assess whether correlational selection generated by pollinators can account for observed phenotypic integration of the two corolla lips in the zygomorphic flowers of the insect‐pollinated Mediterranean shrub, Lavandula latifolia (Labiatae). The experiment consisted of manipulating the upper and lower corolla lips according to a two‐way factorial design with interaction, and involved the complete (lower lip) or extensive (distal two thirds of upper lip) excision of one or the two lips. Per‐flower male (proportion of pollen grains removed) and female (number of pollen tubes down the style) pollination success, and maternal fecundity (seed set), of experimental flowers were evaluated after exposure to natural pollination in the field. Artificially induced corolla variation had no significant effects on either male or female pollination success, and two aberrant floral morphologies, characterized by lack of corolla integration, had a maternal fecundity advantage over normal two‐lipped phenotypes. These results do not support a predominant role of correlational selection by pollinators in maintaining corolla integration in L. latifolia, but are instead consistent with the alternative view that integration is mainly the consequence of genetic and/or developmental factors.     

Covariance and decoupling of floral and vegetative traits in nine Neotropical plants: a re-evaluation of Berg's correlation-pleiades concept

Nearly forty years ago R. L. Berg proposed that plants with specialized pollination ecology evolve genetic and developmental systems that decouple floral morphology from phenotypic variation in vegetative traits. These species evolve separate floral and vegetative trait clusters, or as she termed them, "correlation pleiades." The predictions of this hypothesis have been generally supported, but only a small sample of temperate-zone herb and grass species has been tested. To further evaluate this hypothesis, especially its applicability to plants of other growth forms, we examined the patterns of phenotypic variation and covariation of floral and vegetative traits in nine species of Neotropical plants. We recognized seven specific predictions of Berg's hypothesis. Our results supported some predictions but not others. Species with specialized pollination systems usually had floral traits decoupled (weak correlation; Canna and Eichornia) or buffered (relationship with shallow proportional slope; Calathea and Canna) from variation in vegetative traits. However, the same trend was also observed in three species with unspecialized pollination systems (Echinodorus, Muntingia, and Wedelia). One species with unspecialized pollination (Croton) and one wind-pollinated species (Cyperus) showed no decoupling or buffering, as predicted. While species with specialized pollination usually showed lower coefficients of variation for floral traits than vegetative traits (as predicted), the same was also true of species with unspecialized or wind pollination (unlike our prediction). Species with specialized pollination showed less variation in floral traits than did species with unspecialized or wind pollination, as predicted. However, the same was true of the corresponding vegetative traits, which was unexpected. Also in contrast to our prediction, plants with specialized pollination systems did not exhibit tighter phenotypic integration of floral characters than did species with generalized pollination systems. We conclude that the patterns of morphological integration among floral traits and between floral and vegetative traits tend to be species specific, not easily predicted from pollination ecology, and generally more complicated than R. L. Berg envisaged.     

Trade-off mitigation: a conceptual framework for understanding floral adaptation in multispecies interactions

Explanations of floral adaptation to diverse pollinator faunas have often invoked visitor-mediated trade-offs in which no intermediate, generalized floral phenotype is optimal for pollination success, i.e. fitness valleys are created. In such cases, plant species are expected to specialize on particular groups of flower visitors. Contrary to this expectation, it is commonly observed that flowers interact with various groups of visitors, while at the same time maintaining distinct phenotypes among ecotypes, subspecies, or congeners. This apparent paradox may be due to a gap in our understanding of how visitor-mediated trade-offs could affect floral adaptation. Here we provide a conceptual framework for analysing visitor-mediated trade-offs with the hope of stimulating empirical and theoretical studies to fill this gap. We propose two types of visitor-mediated trade-offs to address negative correlations among fitness contributions of different visitors: visitor-mediated phenotypic trade-offs (phenotypic trade-offs) and visitor-mediated opportunity trade-offs (opportunity trade-offs). Phenotypic trade-offs occur when different groups of visitors impose conflicting selection pressures on a floral trait. By contrast, opportunity trade-offs emerge only when some visitors’ actions (e.g. pollen collection) remove opportunities for fitness contribution by more beneficial visitors. Previous studies have observed disruptive selection due to phenotypic trade-offs less often than expected. In addition to existing explanations, we propose that some flowers have achieved ‘adaptive generalization’ by evolving features to avoid or eliminate the fitness valleys that phenotypic trade-offs tend to produce. The literature suggests a variety of pathways to such ‘trade-off mitigation’. Trade-off mitigation may also evolve as an adaptation to opportunity trade-offs. We argue that active exclusion, or floral specialization, can be viewed as a trade-off mitigation, occurring only when flowers cannot otherwise avoid strong opportunity trade-offs. These considerations suggest that an evolutionary strategy for trade-off mitigation is achieved often by acquiring novel combinations of traits. Thus, phenotypic diversification of flowers through convergent evolution of certain trait combinations may have been enhanced not only through adaptive specialization for particular visitors, but also through adaptive generalization for particular visitor communities. Explorations of how visitor-mediated trade-offs explain the recurrent patterns of floral phenotypes may help reconcile the long-lasting controversy on the validity of pollination syndromes.     

Multiple aspects of the selfing syndrome of the morning glory Ipomoea lacunosa evolved in response to selection: A Qst‐Fst comparison

The frequent transition from outcrossing to selfing in flowering plants is often accompanied by changes in multiple aspects of floral morphology, termed the “selfing syndrome.” While the repeated evolution of these changes suggests a role for natural selection, genetic drift may also be responsible. To determine whether selection or drift shaped different aspects of the pollination syndrome and mating system in the highly selfing morning glory Ipomoea lacunosa, we performed multivariate and univariate Qst‐Fst comparisons using a wide sample of populations of I. lacunosa and its mixed‐mating sister species Ipomoea cordatotriloba. The two species differ in early growth, floral display, inflorescence traits, corolla size, nectar, and pollen number. Our analyses support a role for natural selection driving trait divergence, specifically in corolla size and nectar traits, but not in early growth, display size, inflorescence length, or pollen traits. We also find evidence of selection for reduced herkogamy in I. lacunosa, consistent with selection driving both the transition in mating system and the correlated floral changes. Our research demonstrates that while some aspects of the selfing syndrome evolved in response to selection, others likely evolved due to drift or correlated selection, and the balance between these forces may vary across selfing species.     

Faster clonal turnover in high‐infection habitats provides evidence for parasite‐mediated selection

According to the Red Queen hypothesis for sex, parasite‐mediated selection against common clones counterbalances the reproductive advantage of asexual lineages, which would otherwise outcompete sexual conspecifics. Such selection on the clonal population is expected to lead to a faster clonal turnover in habitats where selection by parasites is stronger. We tested this prediction by comparing the genetic structure of clonal and sexual populations of freshwater snail Potamopyrgus antipodarum between years 2003 and 2007 in three depth‐specific habitats in Lake Alexandrina (South Island, New Zealand). These habitats differ in the risk of infection by castrating trematodes and in the relative proportion of sexual individuals. As predicted, we found that the clonal structure changed significantly in shallow and mid‐water habitats, where prevalence of infection was high, but not in the deep habitat, where parasite prevalence was low. Additionally, we found that both clonal diversity and evenness of the asexual population declined in the shallow habitat. In contrast, the genetic structure (based on F–statistics) of the coexisting sexual population did not change, which suggests that the change in the clonal structure cannot be related to genetic changes in the sexual population. Finally, the frequency of sexuals had no effect on the diversity of the sympatric clonal population. Taken together, our results show a more rapid clonal turnover in high‐infection habitats, which gives support for the Red Queen hypothesis for sex.     

Evolution of the selfing syndrome: Anther orientation and herkogamy together determine reproductive assurance in a self‐compatible plant

Capacity for autonomous self‐fertilization provides reproductive assurance, has evolved repeatedly in the plant kingdom, and typically involves several changes in flower morphology and development (the selfing syndrome). Yet, the relative importance of different traits and trait combinations for efficient selfing and reproductive success in pollinator‐poor environments is poorly known. In a series of experiments, we tested the importance of anther–stigma distance and the less studied trait anther orientation for efficiency of selfing in the perennial herb Arabis alpina. Variation in flower morphology among eight self‐compatible European populations was correlated with efficiency of self‐pollination and with pollen limitation in a common‐garden experiment. To examine whether anther–stigma distance and anther orientation are subject to directional and/or correlational selection, and whether this is because these traits affect pollination success, we planted a segregating F2 population at two native field sites. Selection strongly favored a combination of introrse anthers and reduced anther–stigma distance at a site where pollinator activity was low, and supplemental hand‐pollination demonstrated that this was largely because of their effect on securing self‐pollination. The results suggest that concurrent shifts in more than one trait can be crucial for the evolution of efficient self‐pollination and reproductive assurance in pollinator‐poor habitats.     

Distortion of symmetrical introgression in a hybrid zone: evidence for locus-specific selection and uni-directional range expansion

The fate of species integrity upon natural hybridization depends on the interaction between selection and dispersal. The relative significance of these processes may be studied in the initial phase of contact before selection and gene flow reach equilibrium. Here we study a hybrid zone of two salamander species, Lyciasalamandra antalyana and Lyciasalamandra billae, at the initial phase of hybridization. We quantify the degree and mode of introgression using nuclear and mtDNA markers. The hybrid zone can be characterized as an abrupt transition zone, the central hybrid zone being only c. 400 m, but introgressed genes were traced up to 3 km. Introgression was traced in both sexes but gene flow may be slightly male-biased. Indirect evidence suggests that hybrid males are less viable than females. Introgression occurred at two levels: (1) locus-specific selection led to different allelic introgression patterns independent of species, while (2) asymmetrical species-level introgression occurred predominately from L. antalyana to L. billae due to range expansion of the former. This indicates that foreign genes can be incorporated into novel genomic environments, which in turn may contribute to the great diversity of morphological variants in Lyciasalamandra.     

Paternal‐specific S‐allele transmission in sweet cherry (Prunus avium L.): the potential for sexual selection

Homomorphic self‐incompatibility is a well‐studied example of a physiological process that is thought to increase population diversity and reduce the expression of inbreeding depression. Whereas theoretical models predict the presence of a large number of S‐haplotypes with equal frequencies at equilibrium, unequal allele frequencies have been repeatedly reported and attributed to sampling effects, population structure, demographic perturbation, sheltered deleterious mutations or selection pressure on linked genes. However, it is unclear to what extent unequal segregations are the results of gametophytic or sexual selection. Although these two forces are difficult to disentangle, testing S‐alleles in the offspring of controlled crosses provides an opportunity to separate these two phenomena. In this work, segregation and transmission of S‐alleles have been characterized in progenies of mixed donors and fully compatible pollinations under field conditions in Prunus avium. Seed set patterns and pollen performance have also been characterized. The results reveal paternal‐specific distorted transmission of S‐alleles in most of the crosses. Interestingly, S‐allele segregation within any given paternal or maternal S‐locus was random. Observations on pollen germination, pollen tube growth rate, pollen tube cohort size, seed set dynamics and transmission patterns strongly suggest post‐pollination, prezygotic sexual selection, with male–male competition as the most likely mechanism. According to these results, post‐pollination sexual selection takes precedence over frequency‐dependent selection in explaining unequal S‐haplotype frequencies.     

Geographical and seasonal variation in the intensity of sexual selection in the barn swallow Hirundo rustica: a meta‐analysis

Sexual selection arises from competition among individuals for access to mates, resulting in the evolution of conspicuous sexually selected traits, especially when inter‐sexual competition is mediated by mate choice. Different sexual selection regimes may occur among populations/subspecies within the same species. This is particularly the case when mate choice is based on multiple sexually selected traits. However, empirical evidence supporting this hypothesis at the among‐populations level is scarce. We conducted a meta‐analysis of the intensity of sexual selection on the largest database to date for a single species, the barn swallow (Hirundo rustica), relying on quantitative estimates of sexual selection. The intensity of sexual selection was expressed as the strength (effect size) of the relationships between six plumage ornaments (tail length, tail asymmetry, size of white spots on tail, ventral plumage colour, throat plumage colour and throat patch size) and several fitness proxies related to reproduction, parental care, offspring quality, arrival date from spring migration, and survival. The data were gathered for four geographically separated subspecies (H. r. rustica, H. r. erythrogaster, H. r. gutturalis, H. r. transitiva). The overall mean effect size (Z_r = 0.214; 95% confidence interval = 0.175–0.254; N = 329) was of intermediate magnitude, with intensity of sexual selection being stronger in males than in females. Effect sizes varied during the breeding cycle, being larger before egg deposition, when competition for access to mates reaches its maximum (i.e. in the promiscuous part of the breeding cycle), and decreasing thereafter. In addition, effect sizes from experiments were not significantly larger than those from correlative studies. Finally, sexual selection on different sexually dimorphic traits varied among subspecies. This last result suggests that morphological divergence among populations has partly arisen from divergent sexual selection, which may eventually lead to speciation.