Search

Since the insights by Charles Darwin, heterostyly, a floral polymorphism with morphs bearing stigmas and anthers at reciprocal heights, has become a model system for the study of natural selection. Based on his archetypal heterostylous flower, including regular symmetry, few stamens and a tube, Darwin hypothesised that heterostyly evolved to promote outcrossing through efficient pollen transfer between morphs involving different areas of a pollinator's body, thus proposing his seminal pollination-precision hypothesis. Here we update the number of heterostylous and other style-length polymorphic taxa to 247 genera belonging to 34 families, notably expanding known cases by 20%. Using phylogenetic and comparative analyses across the angiosperms, we show numerous independent origins of style-length polymorphism associated with actinomorphic, tubular flowers with a low number of sex organs, stamens fused to the corolla, and pollination by long-tongued insects. These associations provide support for the Darwinian pollination-precision hypothesis as a basis for convergent evolution of heterostyly across angiosperms., (© 2024. The Author(s).)

Background and Aims: Nectar, a plant reward for pollinators, can be energetically expensive. Hence, a higher investment in nectar production can lead to reduced allocation to other vital functions and/or increased geitonogamous pollination. One possible strategy employed by plants to reduce these costs is to offer variable amounts of nectar among flowers within a plant, to manipulate pollinator behaviour. Using artificial flowers, we tested this hypothesis by examining how pollinator visitation responds to inter- and intra-plant variation in nectar production, assessing how these responses impact the energetic cost per visit., Methods: We conducted a 2 × 2 factorial experiment using artificial flowers, with two levels of nectar investment (high and low sugar concentration) and two degrees of intra-plant variation in nectar concentration (coefficient of variation 0 and 20 %). The experimental plants were exposed to visits (number and type) from a captive Bombus impatiens colony, and we recorded the total visitation rate, distinguishing geitonogamous from exogamous visits. Additionally, we calculated two estimators of the energetic cost per visit and examined whether flowers with higher nectar concentrations (richer flowers) attracted more bumblebees., Key Results: Plants in the variable nectar production treatment (coefficient of variation 20 %) had a greater proportion of flowers visited by pollinators, with higher rates of total, geitonogamous and exogamous visitation, compared with plants with invariable nectar production. When assuming no nectar reabsorption, variable plants incurred a lower cost per visit compared with invariable plants. Moreover, highly rewarding flowers on variable plants had higher rates of pollination visits compared with flowers with few rewards., Conclusions: Intra-plant variation in nectar concentration can represent a mechanism for pollinator manipulation, enabling plants to decrease the energetic costs of the interaction while still ensuring consistent pollinator visitation. However, our findings did not provide support for the hypothesis that intra-plant variation in nectar concentration acts as a mechanism to avoid geitonogamy. Additionally, our results confirmed the hypothesis that increased visitation to variable plants is dependent on the presence of flowers with nectar concentration above the mean., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Spatial (herkogamy) or temporal (dichogamy) separation of sex organs are mechanisms considered to restrict self-pollination and promote outcrossing. Additionally, avoidance of self-interference is proposed to be the driving force for the evolution of these mechanisms, particularly in self-incompatible species. However, species with anthers and stigmas at different levels may increase the rate of imprecise pollen transfer, resulting in pollen discounting. Non-reciprocal stylar dimorphism has been considered a transitional, unstable stage towards the evolution of reciprocal style dimorphism (distyly), to simultaneously avoid interference and lack of precision. In this study we investigate the spatial and temporal separation of sex organs in a population of the style dimorphic and self-incompatible Narcissus broussonetii and their consequences in the reciprocity between the sex organs of morphs and their fecundity. First, we evaluated the relative growth of sex organs after anthesis. Then, we studied the stigma receptivity along the flower lifespan including its effect on seed production in both morphs. Finally, given the weak reciprocity between the sex organs of morphs of this species, we estimated population genetic diversity parameters in Long- and Short-styled plants to explore differences between them as a result of rates of inbreeding due to different mating strategies. We observed that Long-styled plants and Short-styled plants present different strategies to avoid sexual interference and both of them had negative consequences in the reciprocity between the sex organs of morphs. Long-styled plants exhibited a delay in stigma receptivity and a higher growth rate of the style after anthesis, while Short-styled plants presented higher herkogamy and no delay in stigma receptivity. These findings suggest that the avoidance of self-interference, in stylar dimorphic Narcissus species, seems to be more critical than improving of reciprocity between the sex organs of morphs. This might explain why reciprocal herkogamy (distyly) is rare in the genus.

Different pollinators can exert different selective pressures on floral traits, depending on how they fit with flowers, which should be reflected in the patterns of variation and covariation of traits. Surprisingly, empirical evidence in support of this view is scarce. Here, we have studied whether the variation observed in floral phenotypic integration and covariation of traits in Narcissus species is associated with different groups of pollinators. Phenotypic integration was studied in two style dimorphic species, both with dimorphic populations mostly visited by long-tongued pollinators (close fit with flowers), and monomorphic populations visited by short-tongued insects (loose fit). For N. papyraceus, the patterns of variation and correlation among traits involved in different functions (attraction and fit with pollinators, transfer of pollen) were compared within and between population types. The genetic diversity of populations was also studied to control for possible effects on phenotypic variation. In both species, populations with long-tongued pollinators displayed greater phenotypic integration than those with short-tongued pollinators. Also, the correlations among traits involved in the same function were stronger than across functions. Furthermore, traits involved in the transfer of pollen were consistently more correlated and less variable than traits involved in the attraction of insects, and these differences were larger in dimorphic than monomorphic populations. In addition, population genetic parameters did not correlate with phenotypic integration or variation. Altogether, our results support current views of the role of pollinators in the evolution of floral integration., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

Darwin's early work on heterostyly and related style polymorphisms (the presence of two or three style morphs within a population) generated much interest to understand how precise interactions between ecological and genetic mechanisms influence the evolution of floral diversity. Here we tested three key hypotheses proposed to explain the evolution of heterostyly: (i) the presence of self-incompatibility; (ii) the role of pollinators in promoting dissasortative mating; and (iii) floral architecture, which restricts pollinators' movements and ensures more exact pollen deposition on their bodies. We combined data from experiments, field observations, and published studies to test whether evolution of style polymorphism in Narcissus is driven by the incompatibility system, pollinator guilds, or floral architecture, within a phylogenetic framework. Neither differences in pollinator environment nor the presence of genetic self-incompatibility were correlated with presence of style polymorphism. However, our results indicate that the evolution of style polymorphism was driven by the presence of a narrow and long floral tube., (© 2013 The Author(s). Evolution © 2013 The Society for the Study of Evolution.)

Background and Aims: Heterostyly and related style polymorphisms are suitable model systems to evaluate the importance of functional pollinators in the maintenance of population variability. In Narcissus papyraceus different functional pollinators, incompatibility system and flower morphology have been proposed to influence the maintenance of polymorphism through their effect on disassortative mating. Here a test is done to find out if the visitation rate of long- versus short-tongued pollinators correlates with the morph ratio and if the latter is related to other flower traits of the species across its main geographic range., Methods: Floral traits from 34 populations in the south-west of the Iberian Peninsula and in north-west Africa were measured, perianth variation was described and a comparison was made of allometric relationships between sex organs and floral tube. Correlations between pollinator guilds, stigma-anther separation of reciprocal morphs (our proxy for disassortative mating) and morph-ratio variation were analysed. Finally, the incompatibility system of the species in the northern and southern borders of its distribution are described., Key Results: Flowers from southern populations were significantly larger than flowers from centre and northern populations. The abundance of short-styled plants decreased gradually with increasing distance from the core region (the Strait of Gibraltar), with these disappearing only in the northern range. Although there was a significant difference in stigma-anther separation among populations, morph ratio was not associated with reciprocity or floral tube length. Long-style morph frequency increased with short-tongued pollinator visitation rate. Populations from both edges of the distribution range were self-incompatible and within- and between-morph compatible., Conclusions: The style morph ratio changed gradually, whereas perianth trait variation showed abrupt changes with two morphotypes across the range. The positive relationship between the visitation rate of short-tongued pollinators and the decrease of the short-style morph supports our initial hypothesis. The results highlight the importance of different pollinators in determining the presence of style polymorphism.