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Premise: A multi‐omic approach was used to explore proteins and networks hypothetically important for establishing filament dimorphisms in heterostylous Turnera subulata (Sm.) as an exploratory method to identify genes for future empirical research. Methods: Mass spectrometry (MS) was used to identify differentially expressed proteins and differentially phosphorylated peptides in the developing filaments between the L‐ and S‐morphs. RNAseq was used to generate a co‐expression network of the developing filaments, MS data were mapped to the co‐expression network to identify hypothetical relationships between the S‐gene responsible for filament dimorphisms and differentially expressed proteins. Results: Mapping all MS identified proteins to a co‐expression network of the S‐morph's developing filaments identified several clusters containing SPH1 and other differentially expressed or phosphorylated proteins. Co‐expression analysis clustered CDKG2, a protein that induces endoreduplication, and SPH1—suggesting a shared biological function. MS analysis suggests that the protein is present and phosphorylated only in the S‐morph, and thus active only in the S‐morph. A series of CDKG2 regulators, including ATM1, and cell cycle regulators also correlated with the presence of reciprocal herkogamy, supporting our interest in the protein. Conclusions: This work has built a foundation for future empirical work, specifically supporting the role of CDKG2 and ATM1 in promoting filament elongation in response to SPH1 perception. [ABSTRACT FROM AUTHOR]

The R2R3-MYB gene family represents a widely distributed class of plant transcription factors. This gene family plays an important role in many aspects of plant growth and development. However, the characterization of R2R3-MYB genes present in the genome of Coptis teeta has not been reported. Here, we describe the bioinformatic identification and characterization of 88 R2R3-MYB genes in this species, and the identification of members of the R2R3-MYB gene family in species within the order Ranales most closely related to Coptis teeta. The CteR2R3-MYB genes were shown to exhibit a higher degree of conservation compared to those of A. thaliana, as evidenced by phylogeny, conserved motifs, gene structure, and replication event analyses. Cis-acting element analysis confirmed the involvement of CteR2R3-MYB genes in a variety of developmental processes, including growth, cell differentiation, and reproduction mediated by hormone synthesis. In addition, through homology comparisons with the equivalent gene family in A. thaliana, protein regulatory network prediction and transcriptome data analysis of floral organs across three time periods of flower development, 17 candidate genes were shown to exhibit biased expression in two floral phenotypes of C. teeta. This suggests their potential involvement in floral development (anther development) in this species. [ABSTRACT FROM AUTHOR]

Premise: Evolution of cross‐pollination efficiency depends on the genetic variation of flower traits, the pollen vector, and flower trait matching between pollen donors and recipients. Trait matching has been almost unexplored among nonheterostylous species, and we examined whether the match of anther length in pollen donors and stigma length in pollen recipients influences the efficiency of cross‐pollination. To explore potential constraints for evolutionary response, we also quantified genetic variation and covariation among sepal length, petal length and width, stamen length, style length, and herkogamy. Methods: We created 58 experimental arrays of Turnera velutina that varied in the extent of mismatch in the position of anthers and stigmas between single‐flowered plants. Genetic variation and correlations among flower traits were estimated under greenhouse conditions. Results: Style length, but not herkogamy, influenced the efficiency of cross‐pollination. Plants with stamen length that matched the style length of other plants were more efficient pollen donors, whereas those with the style protruding above the stamens of other plants were more efficient pollen recipients. Significant broad‐sense heritability (0.22 > hB2 < 0.42) and moderate genetic correlations (0.33 > r < 0.85) among floral traits were detected. Conclusions: Our results demonstrated that anther‐stigma mismatch between flowers contributed to variation in the efficiency of cross‐pollination. The genetic correlations between stamen length and other floral traits suggests that any change in cross‐pollination efficiency would be driven by changes in style rather than in stamen length. [ABSTRACT FROM AUTHOR]

Herkogamy is the spatial separation of anthers and stigmas within complete flowers, and is a key floral trait that promotes outcrossing in many angiosperms. The degree of separation between pollen-producing anthers and receptive stigmas has been shown to influence rates of selfpollination amongst plants, with a reduction in herkogamy increasing rates of successful selfing in self-compatible species. Self-pollination is becoming a critical issue in horticultural crops grown in environments where biotic pollinators are limited, absent, or difficult to utilise. In these cases, poor pollination results in reduced yield and misshapen fruit. Whilst there is a growing body of work elucidating the genetic basis of floral organ development, the genetic and environmental control points regulating herkogamy are poorly understood. A better understanding of the developmental and regulatory pathways involved in establishing varying degrees of herkogamy is needed to provide insights into the production of flowers more adept at selfing to produce consistent, high-quality fruit. This review presents our current understanding of herkogamy from a genetics and hormonal perspective. [ABSTRACT FROM AUTHOR]

Mammillaria lasiacantha is a small cactus that inhabits at Sierra of Juárez, Chihuahua, Mexico. Their populations have been affected by habitat destruction, illegal extraction, and uncontrolled tourism in the region that steps on the seedlings; therefore, knowing its reproductive biology is necessary for its conservation. Our main questions were: What is the floral morphology and anatomy like? What type of sexual and mating system do M. lasiacantha present? Do the seeds produced by the crosses germinate, how and in what percentage? Sixty individuals from a population of 120 plants were studied at Sierra of Juárez, Juárez Chihuahua from January to August 2021. The study of the floral morpho-anatomy was carried out. Four treatments were applied: natural selfing, forced selfing, geitonogamy and forced outcrossing plus a control. Pollinated flowers, fruits, number of seeds and germination percentage were recorded. Flowers have naked pericarp, funnel-type nectary, nectarostomata, closed style and collateral vascular bundles. The anatomical characteristics of M. lasiacantha agree with those reported for other species of Mammillaria. The flowers of M. lasiacantha present herkogamy, with an obligate outcrossing system. The seeds produced from the outcrossing and control group germinated in 32 and 76%, respectively. The herkogamy and strict outcrossing system of M. lasiacantha should be confirmed in other populations. The seeds are positive photoblastic, without dormancy, but the study of the seed bank and seedling survival in the field would be pertinent. The ex-situ conservation of M. lasiacantha in the site will be feasible, since species high percentage of germination may contribute to produce plants for restoration programs. [ABSTRACT FROM AUTHOR]

Background and Aims Sexual polymorphisms of flowers have traditionally been interpreted as devices that promote cross-pollination, but they may also represent adaptations for exploiting particular pollination niches in local environments. The cross-pollination function of enantiostyly, characterized by flowers having either left- or right-deflected styles, has been uncertain in some lineages, such as the Haemodoraceae, because the positioning of stamens and styles is not always completely reciprocal among morphs. Methods We examined the floral biology of populations of the poorly known species Barberetta aurea (Haemodoraceae) across its native range in South Africa to establish the general features of its enanatiostylous reproductive system and the agents and mechanism of pollen transfer. Results We confirmed that B. aurea has a system of dimorphic enantiostyly. Style morph ratios varied among populations sampled, but with an overall tendency to being equal. Crossing experiments demonstrated that B. aurea is fully self-compatible, that intra- and inter-morph crosses are equally fertile and that it is wholly dependent on pollinator visits for seed production. Pollination is mainly by syrphid flies that transfer the sticky pollen via their wings, which contact the anthers and stigma precisely as they hover during approach and feeding. The majority of syrphid fly visitors feed on a film of highly concentrated nectar situated at the base of ultraviolet-absorbent 'nectar guides'. Because one of the three stamens is deflected in the same direction as the style, we predicted a high likelihood of intra-morph pollination, and this was corroborated by patterns of transfer of coloured dye particles in cage experiments involving syrphid flies. Conclusions Barbaretta aurea exhibits dimorphic enantiostyly and, in contrast to most enantiostylous species, which are pollinated by bees, its flowers are specialized for pollination by syrphid flies. The lack of complete reciprocity of the enantiostylous arrangement of sexual organs facilitates both inter- and intra-morph pollen transfer on the wings of these flies. [ABSTRACT FROM AUTHOR]

Some plants that invade environments outside their native range of distribution often face reproductive challenges such as limited pollinators and mates. Thus, the capacity to reproduce autonomously in the absence of pollinators or mates can facilitate establishment and spread of invasive plants. Here, we study the capacity of Solanum rostratum to reproduce through autonomous self-pollination. Although this plant is self-compatible, it strongly depends on visitation of buzzing bees to reproduce in its native range. We compared floral morphology, pollen: ovule ratios, reproductive phenology, and capacity for autonomous self-pollination of two populations of native distribution (Mexico) and another two populations of the invasive range (USA). We found a reduction in anther-stigma distance, pollen: ovule ratio, and plant height in invasive populations, as well as a shift to increased resource allocation to floral vs. vegetative structures and faster flowering compared to native populations. Importantly, invasive populations show a marked increase in the capacity to produce fruits autonomously (without pollen vectors). Our results suggest that the northern invasion of S. rostratum could have been facilitated by a shift in reproductive strategy from outcrossing to autonomous self-pollination. [ABSTRACT FROM AUTHOR]

Ongoing climatic changes have altered growing conditions for plants by limiting water availability and inducing unprecedented temperature increases, eliciting plant functional responses that compromise floral trait expression, reduce pollen production and promote early self‐pollination. In a controlled greenhouse study, plants of Collinsia heterophylla, an annual mixed‐mated hermaphrodite, were grown under temperature and water stresses. Floral trait responses and related effects on plant reproductive success were recorded. Plants grown under temperature and water stresses were shorter and had fewer leaves at flowering than control plants. Temperature‐stressed plants flowered earlier and had smaller flowers that produced fewer per capita pollen grains than control and water‐stressed plants. While lifetime flower production in plants experiencing temperature stress alone or water stress alone was similar to control plants, those receiving combined temperature and water stresses had significantly lower lifetime flower production. Abiotic stress did not affect investment in female traits such as ovule number per flower, but impacted male traits seen as fewer pollen grains per flower. In plants experiencing abiotic stress, an increase in autonomous self‐pollination was facilitated through a compromise in herkogamy, a sexual interference mechanism that promotes cross‐pollination. Spatial separation between stigma and anther, a measure of herkogamy was zero or negative in stressed plants. Thus, abiotic stress responses in plants did not compromise plant reproductive success as autonomous selfing was augmented. In nature, such responses will likely reduce available nutrition for pollinators through decreased flower and pollen production. [ABSTRACT FROM AUTHOR]

Premise: The relationships between reproductive investment, phenotype, and fitness have been broadly studied in cross‐pollinated plants in contrast to selfing species, which are considered less interesting in this area because they are supposed to be a dead end in any evolutionary pathway. Still, selfing plants are unique systems to study these questions since the position of reproductive structures and traits related to flower size play an important role in female and male pollination success. Methods: Erysimum incanum s.l. is a selfing species complex that has three levels of ploidy (diploids, tetraploids, and hexaploids) and traits that are typically associated with the selfing syndrome. Here, we used 1609 plants belonging to these three ploidies to characterize the floral phenotype and spatial configuration of reproductive structures, reproductive investment (pollen and ovule production), and plant fitness. Then, we used structural equation modelling to analyze the relationship between all these variables across ploidy levels. Results: An increase in ploidy level leads to bigger flowers with anthers exserted farther and more pollen and ovules. In addition, hexaploid plants had higher absolute values for herkogamy, which is positively correlated with fitness. Ovule production significantly mediated the natural selection acting on different phenotypic traits and pollen production, a pattern that is maintained across ploidies. Conclusions: The changes in floral phenotypes, reproductive investment, and fitness with ploidy level suggest that genome duplication can be a driver for transitions in reproductive strategy by modifying the investment in pollen and ovules and linking them with plant phenotype and fitness. [ABSTRACT FROM AUTHOR]

The reproductive strategies of alpine plants are often altered by environmental changes caused by changes in the spatial distribution of the gradient. However, few studies have investigated whether reproductive patterns of the same species vary with elevation. Three natural populations of Primula atrodentata , which are distributed in the eastern Himalayas and have a long flowering period, were selected along the elevation gradients in Shergyla Mountain, Tibet, China. Morph ratio investigation, floral trait measurement, pollinator observation and manipulated pollination experiments were conducted to explore the changes in self-compatibility and floral traits associated with the selfing syndrome along elevation gradients. We found that the breeding system of the S-morph is facultative outcrossing, and that of the L-morph is obligatory outcrossing. We further found that with increasing elevation, the number of pollen and ovules, anther–stigma distance, and inbreeding depression index first increased and then decreased, whereas the seeds per fruit and seed-setting rate under hand self-pollination, pollen limitation and self-incompatibility index tended to decrease first, but then increased. In addition, pollinator diversity and visiting frequency were the highest at the middle elevation (4050 population), which can better explain the nonlinear change in self-fertility with elevation. Our findings provide insights into the evolutionary pattern of self-compatibility in alpine plants along elevational gradients. [ABSTRACT FROM AUTHOR]

Continuous variation in herkogamy has been well reported, however, less attention has been paid to the phenomena that the consecutive expression of two types of herkogamy in the same flower. Euphorbia fischeriana, which have both vertical and lateral herkogamy, show vertical herkogamy during the female phase. However, their gynophores bend to one side with the male phase and show lateral herkogamy. In this study, we observed the effect of successive sexual organs movement on variation in herkogamy traits. By artificially manipulating the flower to present gynophore straightened in the floral center or bend to one side, we attempted to investigate whether herkogamy movement affects pollinator access efficiency, pollen removal and deposition, and seed set ratio. Furthermore, we conducted artificial pollination in the female phase to evaluate the effect of changes in pollination environment on the variations in herkogamy traits. The results showed that gynophore straightened in female phase favors pollen deposition, whereas gynophore bending in male phase was conducive to the removal of pollen. Visitation frequency, pollen deposition and removal, and seed set ratio decreased significantly when the gynophore movement was manipulated. Finally, the bending of gynophore was obviously promoted by pollination. Therefore, the continuous variation of herkogamy in the same flower of E. fischeriana caused by the bending of the gynophore could improve the accuracy of pollination and avoid the interference of the ovary with access efficiency. That may be an adaptive strategy when pollinators are scarce. Furthermore, our study also provides good support for the hypothesis that variations in herkogamy traits are strongly selected by differences in pollination environments. [ABSTRACT FROM AUTHOR]

Seed production on native seed farms has increased to meet the rising demand for plant material for restoration. Although these propagation efforts are necessary for restoration, cultivating wild populations may also result in unintentional selection and elicit evolutionary changes that mimic crop domestication, essentially turning these efforts into artificial domestication experiments. Here, we investigated whether phenotypic and genomic changes associated with domestication occurred in the wildflower Clarkia pulchella Pursh (Onagraceae) by comparing the wild source populations to the farmed population after eight generations of cultivation. At the phenotypic level, the farmed population shifted towards a floral morphology associated with self-pollination, with a significant decrease in both dichogamy and herkogamy. At the genomic level, > 6500 SNPs revealed that mean expected heterozygosity of the farmed population was significantly lower than the wild populations, despite the fact that the farmed population originated from a pool of multiple wild populations. Both the shift towards a selfing phenotype and the loss of diversity are expected consequences of domestication, although the phenotypic shifts in particular occurred much more rapidly than has been observed for other domestication traits. We discuss these results in the context of plant domestication and the implications for retaining the genetic integrity of wild populations during the process of seed production for restoration., (© 2025 The Author(s). Molecular Ecology published by John Wiley & Sons Ltd.)

The evolutionary transition from outcrossing in heterostylous morphs to selfing in homostylous morphs has occurred in many plant lineages. Homostylous variants with a low degree of herkogamy may increase the reproductive advantage of autonomous self‐pollination when there is little inbreeding depression and/or low pollinator visitation. Oxalis corniculata, a self‐compatible perennial herb derived from a tristylous ancestor, is reported to have two floral morphs. Homostyled plants are broadly distributed in Japan, whereas long‐styled plants are reported to occur only in coastal areas. We examined the hypothesis that a reproductive advantage in avoiding inbreeding depression and/or frequent pollinator visits enables long‐styled plants to live in some areas. Studies on floral variation revealed that the degree of herkogamy varied widely and continuously within and among populations. Homostyled and long‐styled morphs were functionally defined based on the ability of autonomous self‐pollination. Homostyled plants were widely distributed, whereas long‐styled plants were found in and around the two coastal populations and relatively inland populations. Pollination experiments provided no evidence of inbreeding depression in either homostyled or long‐styled plants. Pollinator visitation frequency was higher in the two coastal populations than in the other populations. Thus, reproductive disadvantage in long‐styled plants lacking autonomous self‐pollination has reduced in the two coastal populations, and long‐styled plants may be maintained due to a competitive advantage of large seed size. In contrast, we recorded no comparable frequencies of pollinator visitation in other populations containing long‐styled plants, indicating that there are other factors contributing to the maintenance of long‐styled plants in these areas. [ABSTRACT FROM AUTHOR]

Background and aimsThe evolution of selfing from outcrossing may be the most common transition in plant reproductive systems and is associated with a variety of ecological circumstances and life history strategies. The most widely discussed explanation for these associations is the reproductive assurance hypothesis - the proposition that selfing is favoured because it increases female fitness when outcross pollen receipt is limited. Here an alternative explanation, the time limitation hypothesis, is addressed, one scenario of which proposes that selfing may evolve as a correlated response to selection for a faster life cycle in seasonally deteriorating environments.MethodsArtificial selection for faster maturation (early flowering) or for low herkogamy was performed on Clarkia unguiculata (Onagraceae), a largely outcrossing species whose closest relative, C. exilis, has evolved higher levels of autogamous selfing. Direct responses to selection and correlated evolutionary changes in these traits were measured under greenhouse conditions. Direct responses to selection on early flowering and correlated evolutionary changes in the node of the first flower, herkogamy, dichogamy, gas exchange rates and water use efficiency (WUE) were measured under field conditions.Key resultsLines selected for early flowering and for low herkogamy showed consistent, statistically significant responses to direct selection. However, there was little or no evidence of correlated evolutionary changes in flowering date, floral traits, gas exchange rates or WUE.ConclusionsThese results suggest that the maturation rate and mating system have evolved independently in Clarkia and that the time limitation hypothesis does not explain the repeated evolution of selfing in this genus, at least through its indirect selection scenario. They also suggest that the life history and physiological components of drought escape are not genetically correlated in Clarkia, and that differences in gas exchange physiology between C. unguiculata and C. exilis have evolved independently of differences in mating system and life history.

Durian or Durio zibethinus of variety Musang King is growing in popularity and with high international demands. With the ever-increasing demands for fruits, growers are exploring ways to maximize production by looking at the feasibility of planting single or mono varieties in a planting area. Previous investigations revealed that many durian varieties are self-incompatible, and the condition varies from one variety to another. Against this background, the present study evaluated Musang King's compatibility status in fruit sets. The study was conducted in Raub, Pahang, from 2017 through 2018 with five different pollination treatments. Crossing Musang King with D24 showed the highest fruit set rate of 16.28% at harvest and suggested this variety is self-incompatible. Observations on the flowering process revealed that Musang King possessed herkogamy condition, which posed a morphological barrier to self-pollination. The study proposes that Musang King is best planted in a multi-variety planting system instead of mono-variety to achieve a higher rate of fruit sets. [ABSTRACT FROM AUTHOR]

About one-fifth of angiosperms reproduce predominantly through self-fertilization, mediated by morphological and functional flower traits. Autonomous selfing is rare in Melastomataceae, a megadiverse family with most species featuring poricidal anthers and herkogamy. However, previous studies suggested that some Bertolonia spp. were capable of autonomous selfing. Here, we investigated the existence of autonomous selfing in the group and, considering its specialized floral morphology, the mechanisms involved in this process. We conducted studies on floral biology, pollination experiments and pollinator visitation rates for Bertolonia paranaensis (anthers with introrse pores) and Bertolonia mosenii (anthers with extrorse pores). The transient absence of herkogamy and the dynamics of the movements of the flower parts during post-anthesis events promoted delayed selfing in both species. In post-anthesis flowers, pollen grains germinate inside the anthers, the pollen tubes come out through the anther pores and then reach the stigma, penetrating the transmission tissue, and fertilize the ovules. Both species received visits from buzzing bees and have a mixed mating system, i.e. the capacity of cross-pollination and delayed selfing-pollination. We present the first description of an autonomous selfing mechanism for plants with poricidal anthers, for which we coin the name "pollen tube shower". Pollen tube shower promotes reproductive assurance to flowers that have not been pollinated and ensures reproductive success of individuals in the absence of pollinators. [ABSTRACT FROM AUTHOR]

Premise: Widespread associations between selfing rate and floral size within and among taxa suggest that these traits may evolve in concert. Does this association develop immediately because of shared genetic and/or developmental control, or stepwise with selection shaping the evolution of one trait following the other? If the former, then association ought to appear within and across selfing populations. We explore this fundamental question in three populations of the mixed‐mater Collinsia verna where autonomous selfing (AS) ability has been shown to be under selection by the pollination environment. Methods: We grew clonal replicates of C. verna in a controlled environment to characterize broad‐sense genetic correlations among traits within populations and to assess whether divergence in mating system and floral traits among these populations is consistent with their previously observed selection pressures. Results: As predicted by their respective pollination environments, we demonstrate significant genetic divergence among populations in AS ability. However, patterns of divergence in floral traits (petal, stamen, and style size, stigmatic receptivity, and stigma‐anther distance) were not as expected. Within populations, genetic variation in AS appeared largely independent from floral traits, except for a single weak negative association in one population between flower size and AS rate. Conclusions: Together, these results suggest that associations between selfing rate and floral traits across Collinsia species are not reflected at microevolutionary scales. If C. verna were to continue evolving toward the selfing syndrome, floral trait evolution would likely follow stepwise from mating system evolution. [ABSTRACT FROM AUTHOR]

Outcrossing plant species are more likely to exhibit autonomous selfing in marginal habitats to ensure reproduction under conditions of limited pollinator and/or mate availability. Distyly is a classical paradigm that promotes outcrossing; however, little is known about the variation in floral traits associated with distylous syndrome in marginal populations. In this study, we compared the variation in floral traits including stigma and anther height, corolla tube length, herkogamy, and corolla diameter between the central and peripheral populations of the distylous Primula wannanensis, and assessed the variation of floral traits at early and late florescence stages for each population. To evaluate the potential consequences of the variation in floral traits on the mating system, we investigated seed set in each population under both open-pollinated and pollinator-excluded conditions. The flower size of both short- and long-styled morphs was significantly reduced in late-opening flowers compared with early opening flowers in both central and peripheral populations. Sex-organ reciprocity was perfect in early opening flowers; however, it was largely weakened in the late-opening flowers of peripheral populations compared with central populations. Of these flowers, disproportionate change in stigma height (elongated in S-morph and shortened in L-morph) was the main cause of reduced herkogamy, and seed set was fairly high under pollinator-excluded condition. Our results provide empirical support for the hypothesis on the evolution of delayed autonomous selfing in marginal populations of distylous species. Unsatisfactory pollinator service is likely to have promoted reproductive assurance of distylous plants with largely reduced herkogamy mimicking “homostyles.”

Outcrossing plant species are more likely to exhibit autonomous selfing in marginal habitats to ensure reproduction under conditions of limited pollinator and/or mate availability. Distyly is a classical paradigm that promotes outcrossing; however, little is known about the variation in floral traits associated with distylous syndrome in marginal populations. In this study, we compared the variation in floral traits including stigma and anther height, corolla tube length, herkogamy, and corolla diameter between the central and peripheral populations of the distylous Primula wannanensis , and assessed the variation of floral traits at early and late florescence stages for each population. To evaluate the potential consequences of the variation in floral traits on the mating system, we investigated seed set in each population under both open-pollinated and pollinator-excluded conditions. The flower size of both short- and long-styled morphs was significantly reduced in late-opening flowers compared with early opening flowers in both central and peripheral populations. Sex-organ reciprocity was perfect in early opening flowers; however, it was largely weakened in the late-opening flowers of peripheral populations compared with central populations. Of these flowers, disproportionate change in stigma height (elongated in S-morph and shortened in L-morph) was the main cause of reduced herkogamy, and seed set was fairly high under pollinator-excluded condition. Our results provide empirical support for the hypothesis on the evolution of delayed autonomous selfing in marginal populations of distylous species. Unsatisfactory pollinator service is likely to have promoted reproductive assurance of distylous plants with largely reduced herkogamy mimicking "homostyles." [ABSTRACT FROM AUTHOR]

Inter‐morph pollen transfer and its dependence on herkogamy and reciprocity are not completely understood in species with stigma‐height dimorphism. We asked whether total stigmatic pollen loads, inter‐morph fraction of pollen load and reproductive success differed between morphs in Jasminum malabaricum, a species exhibiting stigma‐height dimorphism. We tested whether higher herkogamy and reciprocity resulted in higher inter‐morph pollen deposition and reproductive success.We quantified individual‐level estimates of herkogamy, reciprocity, total stigmatic pollen load, inter‐morph stigmatic pollen fraction and fruit set for both morphs in naturally occurring populations of J. malabaricum.Total pollen load was higher in the long‐styled morph, inter‐morph pollen fraction was higher in the short‐styled morph, but fruit set did not differ between morphs. Higher herkogamy resulted in a higher inter‐morph fraction of pollen load and fruit set in the long‐styled morph of one population. In the other population, only reciprocity was found to be related to inter‐morph pollen deposition.This study is the first to quantify and report natural inter‐morph stigmatic pollen load in a species with stigma‐height dimorphism. Morph‐specific differences in pollen load were similar to patterns commonly observed in heterostylous species. The results highlight the importance of both herkogamy and reciprocity in facilitating inter‐morph pollen transfer. Population‐specific patterns indicate that local environmental factors determine the relative functional importance of herkogamy and reciprocity. [ABSTRACT FROM AUTHOR]

Background and Aims Theory predicts that outcrossing should be more prevalent among perennials than annuals, a pattern confirmed by comparative evidence from diverse angiosperm families. However, intraspecific comparisons between annual and perennial populations are few because such variation is uncommon among flowering plants. Here, we test the hypothesis that perennial populations outcross more than annual populations by investigating Incarvillea sinensis , a wide-ranging insect-pollinated herb native to China. The occurrence of both allopatric and sympatric populations allows us to examine the stability of mating system differences between life histories under varying ecological conditions. Methods We estimated outcrossing rates and biparental inbreeding in 16 allopatric and five sympatric populations in which both life histories coexisted using 20 microsatellite loci. In each population we measured height, branch number, corolla size, tube length and herkogamy for ~30 individuals. In a sympatric population, we recorded daily flower number, pollinator visitation and the fruit and seed set of annual and perennial plants. Key Results As predicted, outcrossing rates (t) were considerably higher in perennial (mean = 0.76) than annual (mean = 0.09) populations. This difference in mating system was also maintained at sympatric sites where plants grew intermixed. In both allopatric and sympatric populations the degree of herkogamy was consistently larger in outcrossing than selfing plants. Perennials were more branched, with more and larger flowers than in annuals. In a sympatric population, annuals had a significantly higher fruit and seed set than perennials. Conclusions Genetically based differences in herkogamy between annuals and perennials appear to play a key role in governing outcrossing rates in populations, regardless of variation in local ecological conditions. The maintenance of mating system and life history trait differentiation between perennial and annual populations of I. sinensis probably results from correlated evolution in response to local environmental conditions. [ABSTRACT FROM AUTHOR]

PREMISE: Flowering plants with poricidal anthers are commonly visited by buzzing bees, which vibrate flowers to extract pollen. However, not all flower visitors are in fact pollinators, and features such as body size and duration of flower visits are important factors in determining pollination effectiveness. We tested whether bee‐to‐flower size relationships predict the pollination effectiveness of flower visitors of a buzz‐pollinated species (Chamaecrista ramosa, Fabaceae). METHODS: We sorted 13 bee taxa into three groups: smaller than, equivalent to ("fit‐size"), and larger than flower herkogamy (spatial separation between anthers and stigma). We expected the latter two groups to touch the stigmas, which would be an indicator of pollination effectiveness, more frequently than the first group. To test this hypothesis, we assessed contact with stigmas, foraging behavior, and duration of visits for the three size groups of bees. RESULTS: Our data reveal that small bees scarcely touched the stigmas, while large and fit‐size bees were the most efficient pollinators, achieving high stigma‐touching rates, conducting much shorter flower visits, and visiting flowers and conspecific plants at high rates during foraging bouts. CONCLUSIONS: The results did not show size‐matching among bees and flowers, as expected, but rather a minimum size threshold of efficient pollinators. The finding of such a threshold is a nonarbitrary approach to predicting pollination effectiveness of visitors to herkogamous flowers with poricidal anthers. [ABSTRACT FROM AUTHOR]

Plant mating system determines, to a great extent, the demographic and genetic properties of populations, hence their potential for adaptive evolution. Variation in plant mating system has been documented between phylogenetically related species as well between populations of a species. A common evolutionary transition, from outcrossing to selfing, is likely to occur under environmental spatial variation in the service of pollinators. Here, we studied two phenotypically (in floral traits) and genetically (in neutral molecular markers) differentiated populations of the annual, insect-pollinated, plant Datura inoxia in Mexico, that differ in the service of pollinators (Mapimí and Cañada Moreno). First, we determined the populations’ parameters of phenotypic in herkogamy, outcrossing and selfing rates with microsatellite loci, and assessed between generation (adults and seedlings) inbreeding, and inbreeding depression. Second, we compared the relationships between parameters in each population. Results point strong differences between populations: plants in Mapimí have, on average, approach herkogamy, higher outcrossing rate (tm = 0.68), lower primary selfing rate (r = 0.35), and lower inbreeding at equilibrium (Fe = 0.24) and higher inbreeding depression (δ = 0.25), than the populations of Cañada. Outcrossing seems to be favored in Mapimí while selfing in Cañada. The relationship between r and Fe were negatively associated with herkogamy in Mapimí; here, progenies derived from plants with no herkogamy or reverse herkogamy had higher selfing rate and inbreeding coefficient than plants with approach herkogamy. The difference Fe–F is positively related to primary selfing rate (r) only in Cañada Moreno which suggests inbreeding depression in selfing individuals and then genetic purging. In conclusion, mating system evolution may occur differentially among maternal lineages within populations of Datura inoxia, in which approach herkogamy favors higher outcrossing rates and low levels of inbreeding and inbreeding depression, while no herkogamy or reverse herkogamy lead to the evolution of the “selfing syndrome” following the purge of deleterious alleles despite high inbreeding among individuals.

Plant mating system determines, to a great extent, the demographic and genetic properties of populations, hence their potential for adaptive evolution. Variation in plant mating system has been documented between phylogenetically related species as well between populations of a species. A common evolutionary transition, from outcrossing to selfing, is likely to occur under environmental spatial variation in the service of pollinators. Here, we studied two phenotypically (in floral traits) and genetically (in neutral molecular markers) differentiated populations of the annual, insect-pollinated, plant Datura inoxia in Mexico, that differ in the service of pollinators (Mapimí and Cañada Moreno). First, we determined the populations' parameters of phenotypic in herkogamy, outcrossing and selfing rates with microsatellite loci, and assessed between generation (adults and seedlings) inbreeding, and inbreeding depression. Second, we compared the relationships between parameters in each population. Results point strong differences between populations: plants in Mapimí have, on average, approach herkogamy, higher outcrossing rate (tm = 0.68), lower primary selfing rate (r = 0.35), and lower inbreeding at equilibrium (Fe = 0.24) and higher inbreeding depression (d = 0.25), than the populations of Cañada. Outcrossing seems to be favored in Mapimí while selfing in Cañada. The relationship between r and Fe were negatively associated with herkogamy in Mapimí; here, progenies derived from plants with no herkogamy or reverse herkogamy had higher selfing rate and inbreeding coefficient than plants with approach herkogamy. The difference Fe-F is positively related to primary selfing rate (r) only in Cañada Moreno which suggests inbreeding depression in selfing individuals and then genetic purging. In conclusion, mating system evolution may occur differentially among maternal lineages within populations of Datura inoxia, in which approach herkogamy favors higher outcrossing rates and low levels of inbreeding and inbreeding depression, while no herkogamy or reverse herkogamy lead to the evolution of the "selfing syndrome" following the purge of deleterious alleles despite high inbreeding among individuals. [ABSTRACT FROM AUTHOR]

Premise of research. Analyses of the evolution of floral morphology and symmetry broaden our understanding of the drivers of angiosperm diversification. Integrated within a flower, labile floral characters produce different phenotypes that promote variable interactions with pollinators. Thus, investigation of floral evolution may help with inferring potential historic transitions in pollinator modes and ecological pressures that generated present diversity. This study aims to explore the morphological evolution of flowers in Miconieae, a species-rich Neotropical tribe within the Melastomataceae. Despite a constrained floral plan, Melastomataceae manage to achieve a variety of floral phenotypes appealing to diverse pollinator types, with a majority of the species requiring specialized "buzz pollination" by bees. However, previous research of Miconieae documented several instances of convergent evolution of phenotypes associated with generalized pollination strategies. We explore the floral morphological evolution of Miconieae in a phylogenetic context to understand how diversification relates to different phenotypes and how common evolution of generalized pollination systems is within this tribe. Methodology. We constructed the largest species-level phylogeny of the Miconieae and combined it with morphological data on a variety of floral characteristics for more than 350 species scored from field photographs. Trait evolution was then analyzed using ancestral state reconstruction with stochastic character mapping. Trait correlation was estimated with Pagel's statistical method for discrete characters and with regression analysis of phylogenetically independent contrasts of continuous characters. We analyzed diversification in the tribe with Bayesian analysis of macroevolutionary mixtures and explored the effect of character state evolution on diversification with the binary state speciation and extinction approach. Pivotal results. Our analyses reveal rampant convergent and correlated evolution of multiple characters indicative of pollinator-mediated selective pressures. Conclusions. We confirm several parallel trends in the evolution of generalized floral phenotypes and find an association between generalization trends and increased diversification rates that may be related to colonization of highland environments. [ABSTRACT FROM AUTHOR]

Background and Aims Herkogamy, or anther–stigma separation, is known to reduce self-pollen deposition, but little is known about the relative efficacy of different modes or conformations of herkogamy. We assessed the effectiveness of vertical versus lateral herkogamy in preventing or promoting self-pollen deposition in the annual herb Lysimachia arvensis , a plant with lineages that differ in flower colour, and in which flowers first display lateral and then vertical herkogamy. Because mating between the two lineages compromises fitness through the production of low-quality hybrid offspring, we tested the prediction that individuals sampled from sites occupied by both lineages should have flowers that promote autonomous self-pollen deposition and self-fertilization as a result of selection to reduce deleterious reproductive interference. Methods We characterized variation in herkogamy within and among 25 pure and mixed populations of L. arvensis in its European range and assessed the effectiveness of lateral versus vertical herkogamy in avoiding self-pollen deposition. Results Lateral herkogamy was more effective than vertical herkogamy in limiting self-pollen deposition. In the case of vertical herkogamy, only approach herkogamy was effective. Lineages showed consistent differences in herkogamy traits. In general, angles were smaller for blue than red flowers in most populations, and blue flowers showed approach herkogamy, while red flowers showed predominantly reverse herkogamy. In sympatry, the red lineage showed a reduction of both herkogamy traits while for the blue lineage only lateral herkogamy was reduced. Conclusions Our results demonstrate that pollen deposition is affected not only by the degree but also the spatial conformation of herkogamy. They also highlight reduced herkogamy as a potential mechanism for promoting reproductive assurance under pollen limitation, as well as for avoiding reproductive interference between genetically divergent lineages. [ABSTRACT FROM AUTHOR]

The location of nectar secretion in flowers of Jaltomata has not been identified with certainty until now: removal of the corolla and androecium from one side of living flowers allowed us to see, in progress, nectar secretion by the ovarian nectary. We studied Jaltomata quipuscoae, a wild plant that grows in southern Peru and produces copious, red floral nectar. Unmanipulated flowers do not set fruit in a pollinator-free greenhouse, demonstrating lack of autogamy, but self-compatibility was demonstrated by manual self-pollinations leading to fruit-set. Anther dehiscence is staggered with the anthers of a flower dehiscing over hours on the same day. The corolla and nectar are UV-absorptive. Flowers last 4–10 days, are usually protogynous during the first day the corolla is open, and do not close for the night. [ABSTRACT FROM AUTHOR]

Introduced plants become decoupled from their usual pollinators and by relying on resident pollinator species or other reproductive strategies become established in new areas, spread and, eventually, invade. Here, using breeding system experiments, we studied the reproductive biology of the bird-pollinated South American species Nicotiana glauca across native and non-native areas, both inside and outside the range of flower-visiting birds. In the native range, where the species is visited by hummingbirds, open cross-pollinated flowers set as many seeds and almost as many fruits as open pollination controls, suggesting that pollinators make a major contribution to reproductive success. In South Africa, the flowers were pollinated by sunbirds which, although less efficiently, also contributed to N. glauca reproduction, replacing hummingbirds. In contrast, in Mallorca, where nectar feeding birds are absent, fruit production in open cross-pollinated flowers was near zero, and significantly lower than in open pollination controls, suggesting that reproduction is almost entirely by autonomous self-pollination. Hand-pollination experiments showed that the species is self-compatible throughout its range. Pollinator exclusion experiment showed that N. glauca relies on self-pollination only in pollinator poor areas, where plants have a much higher capacity for autonomous self-pollination than elsewhere. A reduction in anther-stigma distance does not seem to account the higher self-pollination capacity in the non-native environment without pollinators. Despite probable evolutionary adaptation, and flexibility in pollinator association and mating system, seed production in the introduced range was somewhat pollen limited, suggesting that the mismatch between N. glauca and its novel pollination environment might retard invasion. [ABSTRACT FROM AUTHOR]

Climate change can negatively impact plant–pollinator interactions, and reduce outcross pollination. For reproductive assurance, an increased capacity for autonomous selfing should benefit the persistence of plants under new temperature conditions. Plastic responses of the autonomous selfing capacity to climate change may occur indirectly due to changes in floral traits associated with this capacity. We tested whether the mixed mating plant Mimulus guttatus is capable of plastic changes in floral traits favoring autonomous selfing in response to temperature changes. In seven growth chambers, we grew M. guttatus originating from a large range of latitudes (from 37.89° N to 49.95° N) and thus home temperatures in North America, and experimentally assessed the (autonomous) selfing and outcrossing capacities of the plants. With an increase in the difference between the overall mean daytime and nighttime experimental test temperature and home temperature, flower length and width decreased. The plastic response in flower size suggests that plants may be more successful at autonomous selfing. However, we did not find direct evidence that M. guttatus responded to increased temperature by an increased autonomous selfing capacity. With an increase in temperature difference, the odds of seed production, number of seeds, and individual seed mass decreased. Our results indicate that global warming and the associated increase in extreme temperature events may be detrimental to the reproduction and thus persistence of some plants. [ABSTRACT FROM AUTHOR]

Background and Aims To predict the evolutionary consequences of pollinator declines, we need to understand the evolution of delayed autonomous self-pollination, which is expected to evolve as a mechanism of reproductive assurance when cross-pollination becomes unreliable. This involves estimating the costs of increased levels of selfing as well as those associated with floral senescence. Methods We studied the mechanisms and costs of delayed self-pollination in the mixed-mating vine Dalechampia scandens (Euphorbiaceae) by first assessing among-population variation in herkogamy and dichogamy, which together determine the rate and timing of autonomous self-pollination. We then tested whether floral longevity responds plastically to delayed pollination. Finally, we assessed the costs of delayed self-pollination in terms of seed number and size, explicitly separating inbreeding depression from effects of floral senescence. Key Results Herkogamy varied extensively, while variation in dichogamy was more limited. Unpollinated blossoms increased their longevity, but seed quantity and quality decreased with increasing delays in pollination, independently of inbreeding depression. Conclusions In D. scandens , earlier autonomous selfing is facilitated by reduced herkogamy rather than reduced protogyny, providing reproductive assurance while maintaining the possibility for outcrossing events. Effective early autonomous self-pollination may evolve under reduced cross-pollination reliability in response to costs associated with floral senescence. [ABSTRACT FROM AUTHOR]

Premise: Distyly, a plant breeding system characterized by two floral morphs that have reciprocal positioning of anthers and stigmas, is known from at least 27 angiosperm families, making it an excellent example of convergent evolution. The various manners in which patterns of floral development produce the distinct anther and stigma heights in each morph remain largely unexplored from developmental and evolutionary perspectives. Methods: In 15 species representing at least 12 origins of distyly, heights and lengths of floral organs in each morph throughout development were examined using light microscopy. Patterns of floral organ development were determined and compared among species. Family‐level phylogenies of distylous species and relatives were reconstructed, and patterns of ancestral herkogamy were resolved. Results: Differences in floral development between morphs resulted in 12 patterns leading to the anther and stigma positions characterizing distyly. Distylous species evolved from ancestors with different types of herkogamy, with approach herkogamy and lack of herkogamy resolved most frequently. Conclusions: Seven of the 12 patterns of floral development are known from only one species, with three other patterns described among pairs of close relatives. The most common pattern of floral development, described from at least seven genera, involves for anther heights, distinct intermorph growth rates and for stigma heights, growth rates that differ between morphs only during later development. This pattern is common among subclass Lamiidae, suggesting canalized development within the taxon. Among distylous species, the same type of ancestral herkogamy can give rise to different patterns of floral development. [ABSTRACT FROM AUTHOR]

In flowering plants, the shift from outcrossing to selfing is associated with a set of correlated changes in morphological and reproductive features known as the 'selfing syndrome'. Species of the sub-endemic Mediterranean genus Centaurium Hill (Gentianaceae) exhibit a wide array of flower traits related to pollination biology and different ploidy levels. We explored if the evolutionary transitions of seven flower traits and life cycle, typically associated with the selfing syndrome, are related to polyploidy, diversification patterns, divergence times and the geological and climatic history of the Mediterranean Basin. Using 26 species of Centaurium we reconstructed a phylogenetic tree, inferred the ancestral states of the selected traits, estimated their phylogenetic signal and tested the correlative evolution among them. We found a significant increase in diversification rates during the Quaternary. Anther length, flower size, herkogamy and polyploidy undergo rapid state transitions without phylogenetic signal that could be the result of adaptation for selfing. Changes in character states do not show evidence of correlative evolution among them, as would be predicted during the evolution of selfing syndrome. The evolution of reproductive traits in Centaurium has probably relied on a more diverse array of drivers than just reproductive assurance or polyploidy events. [ABSTRACT FROM AUTHOR]

Most heterostylous species show self- and intra-morph incompatibility and models established for such taxa have traditionally been applied to the evolution of stylar dimorphism and heterostyly in Narcissus, a genus with late-acting self-incompatibility. The model of Lloyd and Webb (1992a,b) proposed that, in an approach-herkogamous ancestor, stylar dimorphism and heterostyly appeared consecutively as a result of two single mutations selected positively to enhance cross-pollination. Most polymorphic Narcissus are stylar dimorphic with two anther whorls, the lower positioned in the middle of a narrow floral tube, and style lengths that locate the stigmas above or below the low-level anthers. Here, I propose that in an ancestor with open-tubed flowers, late-acting self-incompatibility and variable style length, the narrowing of the floral tube increased self-pollination and ovule discounting in individuals with the stigma at the same height as the low-level anthers, imposing gradual disruptive selection against this phenotype and causing the bimodal distribution of style lengths. This hypothesis stresses the need of avoiding self-interference for the selection of stylar dimorphism, but excludes neither the promotion of cross-pollination as a force for subsequent evolutionary steps towards heterostyly in the genus nor the need of inter-morph pollination for the maintenance of polymorphism.

Background: Hybridization is a key driver of plant speciation, and understanding ongoing hybridization provides insights into the underlying process. We studied a hybrid complex in Mediterranean Cyclamen between common C. repandum and rare C. balearicum on Sardinia. Aims: We analyzed variation in the frequency and morphology of different floral types to assess the degree of hybridization and analyze the ecological niche to better understand factors that influence the maintenance of hybrid populations. Methods: We recorded the frequency of the different flower color-types in 45 populations and measured floral traits in 10 of them. We characterized the spatial extent and climatic niche of this hybrid complex. We analyzed floral type variation in four traits (flower size, style and anther length, herkogamy). Results: Hybrid populations occupy a wide area on the NW coast of the island (~1000km2). Sites dominated by hybrid floral types are ecologically marginal for the common parent. The C. repandum-like type is present in all populations and shows a decrease in style and anther length and herkogamy in populations containing a greater proportion of hybrids. Conclusions: This hybrid complex provides an ideal situation to examine reproductive isolation due to variation in the ecological niche of different floral types. [ABSTRACT FROM AUTHOR]

Premise of the Study: Turnover in biotic communities across heterogeneous landscapes is expected to lead to variation in interactions among plants, their mutualists, and their antagonists. Across a fragmented landscape in northern Costa Rica, populations of the euphorb vine Dalechampia scandens vary widely in mating systems and associated blossom traits. Previous work suggested that populations are well adapted to the local reliability of pollination by apid and megachilid bees. We tested whether variation in the intensity of predispersal seed predation by seed weevils in the genus Nanobaris also contributes to the observed variation in blossom traits. Methods: We studied spatiotemporal variation in the relationships between floral advertisement and the probability of seed predation within three focal populations. Then we assessed among‐population covariation of predation rate, pollination reliability, mating system, and blossom traits across 20 populations. Key Results: The probability of seed predation was largely unrelated to variation in floral advertisement both within focal populations and among the larger sample of populations. The rate of seed predation was only weakly associated with the rate of cross‐pollination (allogamy) in each population but tended to be proportionally greater in populations experiencing less reliable pollination. Conclusions: These results suggest that geographic variation in the intensity of antagonistic interactions have had only minor modifying effects on the evolutionary trajectories of floral advertisement in plant populations in this system. Thus, pollinator‐driven floral trait evolution in D. scandens in the study area appears not to be influenced by conflicting seed‐predator‐mediated selection. [ABSTRACT FROM AUTHOR]

Premise of the Study: Nursery pollination systems can range from obligate to facultative. In a system where generalists provide substantial pollination service, an important question is whether the cost of seed predation outweighs the benefit provided by the nursery pollinator to cause the plant to evolve toward more generalized pollination. Using a facultative system native to North America, we tested whether nursery pollinator vs. strictly mutualistic generalists affect mating‐system parameters of the host plant and explored the implications for long‐term coevolution. Methods: We used paternity analyses with 11 microsatellite markers to characterize the mating system of Silene stellata when pollination service is primarily through the nursery pollinator Hadena ectypa and generalist moths. Key Results: Our experimental population of S. stellata was predominantly outcrossing (average outcrossing rate t = 0.83), and mating‐system parameters were similar between pollinator groups. We detected significant correlations in both selfing and outcrossed paternity at the fruit and maternal family level, corresponding to limited pollen dispersal (mean = 3.9 m). Among individuals, variation in anther–stigma separation was positively associated with outcrossing rate, which suggests the importance of herkogamy in preventing selfing. Conclusions: Correlated paternity suggests that seeds from the same fruit and/or plants are sired by a limited number of pollen donors, resulting from low pollen dispersal and potential male–male competition. The similar mating‐system parameters of the two pollinator groups suggest that selection for higher outcrossing in S. stellata is likely to be through floral design rather than through increased pollinator specialization with H. ectypa. [ABSTRACT FROM AUTHOR]

Heterostyly, i.e. the reciprocal positioning of anthers and stigmas, is a floral polymorphism that is thought to promote disassortative (i.e. between‐morph) pollination and to maintain genetic diversity within populations. Recent research, however, has shown several cases of heterostylous plant species in which the reciprocal positioning of the sexual organs varies, which may affect the likelihood of ‘legitimate’ pollination between compatible morphs, and hence morph fitness and ultimately the stability of this floral polymorphism. In this study, we investigated floral morphology and stigmatic pollen deposition patterns in nine Pulmonaria species (Boraginaceae) across Europe to test the hypothesis that sexual organ reciprocity is related to legitimate pollen deposition. We used a recently developed adaptive accuracy concept to assess the level of reciprocity and to relate inaccuracies in positioning of sexual organs to legitimate pollen deposition measured on stigmas in the field. The nine investigated Pulmonaria species showed substantial deviations from perfect reciprocity, with total inaccuracies varying between 2.9 and 20.3% relative to the squared mean of all anther and stigma heights in the population. In almost all cases, most of the inaccuracy was generated by the high‐level organs. Total pollen deposition was always higher on stigmas of long‐styled individuals, but stigmas of short‐styled individuals received significantly more legitimate pollen than stigmas of long‐styled individuals (71.23% and 38.75% respectively). We also found a weak but significant relationship between measures of inaccuracy and the level of disassortative pollination. Under these conditions of pollen flow, the distylous mating system is predicted to persist in only eleven (27%) of all sampled populations. Overall, these results indicate that deviations from perfect reciprocal herkogamy are common in Pulmonaria and have an impact on legitimate pollen deposition. [ABSTRACT FROM AUTHOR]

The benefits of self-fertilization can vary across environments, leading to selection for different reproductive strategies and influencing the evolution of floral traits. Although stressful conditions have been suggested to favour self-pollination, the role of climate as a driver of mating-system variation is generally not well understood. Here, we investigate the contributions of local climate to intraspecific differences in mating-system traits in Clarkia pulchella Pursh in a common-garden growth chamber experiment. We also tested for plastic responses to soil moisture with watering treatments. Herkogamy (anther-stigma spacing) correlated positively with dichogamy (timing of anther-stigma receptivity) and date of first flower, and northern populations had smaller petals and flowered earlier in response to experimental drought. Watering treatment alone had little effect on traits, and dichogamy unexpectedly decreased with annual precipitation. Populations also differed in phenological response to watering treatment, based on precipitation and winter temperature of their origin, indicating that populations from cool and dry sites have greater plasticity under different levels of moisture stress. While some variation in floral traits is attributable to climate, further investigation into variation in pollinator communities and the indirect effects of climate on mating system can improve our understanding of the evolution of plant mating. [ABSTRACT FROM AUTHOR]

Numerous bladderwort (Utricularia) species are distributed worldwide, but their reproductive biology is rarely investigated. Bladderworts are known to depend on tiny organisms to meet a significant proportion of their energy requirement by trapping them in bladders. However, information on the extent of their reliance on insects for pollination success is limited. We examined the reproductive strategy of two Utricularia species viz. Utricularia praeterita and U. babui, endemic to Western Ghats, India. The main aspects of the investigation involved floral biology, breeding system, pollination mechanism, and reproductive success. Flowers of both the species are structured for outbreeding through entomophilous floral suites, herkogamy, protandrous dichogamy and sensitive lobes of the stigma. With nearly 65% natural fruit-set, both the species appeared to be sufficiently open-pollinated. However, pollinators failed to show in plants of U. praeterita while in U. babui there was an apparent mismatch between the extent of fruit-set and pollinator visits. The study demonstrated that in the absence/insufficient visits of pollinators, the two species resort to autonomous selfing. In U. babui, denser patches of plants appeared to be crucial for attracting the pollinators. Both species are self-compatible, and reproductive success is predominantly achieved by delayed autonomous selfing. The sensitive stigma in the species fails to prevent selfing due to diminished herkogamy during the late anthetic stages. It is inferred that in the pollinator-limited environment, delayed selfing contributes to absolute natural fecundity in U. praeterita, while it produces a mixed progeny in U. babui. [ABSTRACT FROM AUTHOR]

Abstract: Mixed mating and variation in outcrossing rate among populations of the same species are common. Outcrossing can be affected by pollinator activity and floral traits that facilitate or impede autonomous self‐fertilization. However, the relative contribution of pollen limitation and evolved differences in the ability to self‐fertilize to variation in the mating system is poorly understood and can only be disentangled using an experimental approach. We placed arrays of plants from eight Campanula americana populations that varied in autonomous selfing ability (hereafter “autonomy”) in sun and shade habitats to create high‐ and low‐pollination environments. Floral visitors were observed and pollen limitation, dichogamy, herkogamy and pollen persistence within the flower were measured. Outcrossing rate was estimated for a subset of the arrays. Pollen limitation was over three times higher in the shade than the sun due to differences in pollinator visitation. Populations with high levels of autonomy displayed greater pollen persistence and dichogamy in the more pollen‐limited environments than in the high‐pollination environments. In these high‐autonomy populations, outcrossing rates were tightly associated with pollen limitation, with reduced outcrossing under strong pollen limitation. In contrast, populations with intermediate autonomy levels had less plastic floral traits and their outcrossing rates changed little with pollen limitation. Mating system was shaped by both the pollination environment and the magnitude of plasticity in floral traits. The experimental approach used here revealed that plasticity in floral traits provides the ability to adjust mating system in response to limited potential for outcrossing. The lack of plasticity in some populations could explain mismatches between pollinator activity and mating system in natural populations. Finally, flexibility in the mating system may help explain the prevalence of mixed mating. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

Background: Autonomous selfing can be favoured by reduced floral herkogamy, dichogamy and pollinator visitation. Autonomous selfing diminishes as pollinator abundance increases; however, the ways pollinators contribute to such result have not been tested. Pollinators can reduce the occurrence of autonomous selfing by two not mutually exclusive mechanisms: successful pollen deposition on stigmas and pollen removal from anthers. Aims: We tested the role of herkogamy and protandry on autonomous selfing and the role of pollinators to reduce it via pollen removal. Methods: We studied Lepechinia floribunda (Lamiaceae) in two natural populations in the Chaco Serrano forest, Argentina. Pollination treatments were conducted to test if floral herkogamy and pollen removal by one pollinator visitation decreased seed set by autonomous selfing. Results: Seed set in pollinator exclusion and emasculation treatments were higher in flowers with lack of herkogamy and selfing increased seed set only in these flowers. Pollen removal during the male phase decreased by 72% the probability of autonomous self-pollination. Conclusions: Flowers with lack of herkogamy are reproductively more advantageous than flowers with approach herkogamy independently of pollinator abundance. We demonstrated for the first time that only one pollinator visitation during the male phase can strongly decrease autonomous selfing by pollen removal. [ABSTRACT FROM AUTHOR]

Abstract: Reproductive success of a plant species is largely influenced by the outcome of mating pattern in a population. It is believed that a significantly larger proportion of animal‐pollinated plants have evolved a mixed‐mating strategy, the extent of which may vary among species. It is thus pertinent to investigate the key contributors to mating success, especially to identify the reproductive constraints in depauperate populations of threatened plant species. We examined the contribution of floral architecture, pollination mechanism and breeding system on the extent of outcrossing rate in a near‐threatened tree species, Wrightia tomentosa. The breeding system was ascertained from controlled pollination experiments. In order to determine outcrossing rate, 60 open‐pollinated progeny were analysed using an AFLP markers. Although the trees are self‐compatible, herkogamy and compartmentalisation of pollen and nectar in different chambers of the floral tube effectively prevent spontaneous autogamy. Pollination is achieved through specialised interaction with moths. Differential foraging behaviour of settling moths and hawkmoths leads to different proportions of geitonogamous and xenogamous pollen on the stigma. However, most open‐pollinated progeny were the result of xenogamy (outcrossing rate, tm = 0.68). The study shows that floral contrivances and pollination system have a strong influence on mating pattern. The differential foraging behaviour of the pollinators causes deposition of a mixture of self‐ and cross‐pollen to produce a mixed brood. Inbreeding depression and geitonogamy appear to play a significant role in sustaining mixed mating in this species. [ABSTRACT FROM AUTHOR]

Premise of Study: In a seminal body of theory, Lloyd showed that the fitness consequences of selfing will depend on its timing in anthesis. Selfing that occurs after opportunities for outcrossing or pollen dispersal can provide reproductive assurance when pollinators are limited and is expected to incur little cost, even when inbreeding depression is high. As a result, delayed selfing is often interpreted as a “best‐of‐both‐worlds” mating system that combines the advantages of selfing and outcrossing. Methods: We surveyed 65 empirical studies of delayed selfing, recording floral mechanisms and examining information on inbreeding depression, autofertility, and other parameters to test the support for delayed selfing as a best‐of‐both‐worlds strategy. Key Results: Phylogenetic distribution of the diverse floral mechanisms suggests that some basic floral structures may predispose plant taxa to evolve delayed selfing. Delayed selfing appears to serve as a best‐of‐both‐worlds strategy in some but not all species. While the capacity for autonomous selfing is often high, it is lower, in some cases, than in related species with earlier modes of selfing. In other delayed‐selfers, low inbreeding depression and reduced investment in corollas and pollen suggest limited benefits from outcrossing. Conclusions: Despite a growing literature on the subject, experimental evidence for delayed selfing is limited and major gaps in knowledge remain, particularly with respect to the stability of delayed selfing and the conditions that may favor transitions between delayed and earlier selfing. Finally, we suggest a potential role of delayed selfing in facilitating transitions from self‐incompatibility to selfing. [ABSTRACT FROM AUTHOR]

In plants capable of clonal reproduction, demographic and ecological investigations are essential for understanding the factors contributing to population diversity. The perennial, clonal orchid Cypripedium calceolus has a variable status in Europe and Asia, being threatened with extinction in some countries and abundant in others. In Estonia, thriving populations can be observed in different biogeographical regions. The aim of this study was to characterize genetic diversity and differentiation in Estonian populations, by comparing the current variation with long-term observations of the clonal system and taking into account climatic variables. Diversity at 11 microsatellite loci for 11 populations representing the boreal and nemoral regions was investigated. Some signatures of bottleneck and recent founder events were detected, with a high genetic diversity in terms of heterozygosity and allele diversity and a pattern of differentiation not strictly correlated with geographical factors. Globally, information regarding population dynamics in C. calceolus provides the possibility to inform conservation management, prevent loss of ancestral variation and promote intra-population divergence through the combination of gene flow and ecological change related to the clonal system. [ABSTRACT FROM AUTHOR]

Background: Heterostyly is a genetic polymorphism characterized by a system of sporophyte auto-incompatibility in which the plant populations show two (distyly) or three (tristyly) floral morphs. This reproductive system is known as a promoter of cross-pollination, assuming similar reproductive success between morphs. Hyphothesis: For this assumption to take place, the pollinators must respond in a similar manner to attraction floral traits in both morphs, maintaining a symmetric flow of pollen. We hypothesize that floral damage by antagonists should correspond to similarities or differences in atractiveness among floral morphs. Studied species: We tested this hypothesis in the distylous shrub Bouvardia ternifolia (Rubiaceae). Methods: we evaluated the magnitude of the nectar theft, foliar herbivory, and seed predation by assessing individual plants of both floral morphs throughout their flowering period (June-July 2015) under natural conditions. At the end of the flowering season, we quantified fruit and seed production. Results: The intensity of the herbivore and nectar thieves in this distylous population is similar between morphs, as well as the size of their floral displays. The number of seeds and fruits produced was also similar between morphs. Conclusions: Our study showed that selection pressures imposed by antagonists were similar to both floral morphs in the studied population of Bouvardia ternifolia.