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PREMISE OF THE STUDY 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/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. KEY 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. This article is protected by copyright. All rights reserved.

The dependence of commercial crops on pollinators is closely related to their reproductive strategies and has direct impacts on production yields. Fruit production of Averrhoa carambola, the star fruit, is highly dependent on bee pollination. This distylous species may exhibit monomorphism associated with self-compatibility, a condition that may influence its degree of dependence on pollinators for fruit set. We evaluated the floral morphology and mating system of A. carambola and their relationships with pollinator dependence in an apparently pin-monomorphic orchard to address the following questions: Is the orchard indeed pin-monomorphic? Is the orchard self-compatible? Are pollination services sufficient to reach maximum productivity? Is fruit set related to the number of visits? We investigated floral morphometrics, mating system, and pollination following standard methods of plant reproduction research. The orchard exhibited pin-monomorphism and self-compatibility. Herkogamy strongly impaired spontaneous self-pollination, emphasizing the relevance of pollinators to crop yield. Considering that Apis mellifera was the only pollinator observed, and that the orchard showed pollen limitation, we recommend the implementation of pollinator management strategies. Additional studies will be needed to assess if the atypical distyly observed here occurs in other situations.

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.

Intraspecific floral variations may affect the degree of generalization and efficacy of pollinators, with consequences for plant reproductive success. Such effects could be greater for heterostylous plants because morphological variation can alter reciprocity between morphs. We evaluated the frequency, composition, and efficacy of pollinators, and the reciprocity (measured by population inaccuracy) of Psychotria nuda, a distylous species of Rubiaceae, in a montane forest. Moreover, we assessed the effects of corolla length on reciprocity and reproductive success. We recorded eighteen species of floral visitors, belonging to four functional groups. The frequency of visits differed among groups with butterflies being the most frequent visitors, in contrast to hummingbird-pollinated lowland populations. This difference did not affect reproductive success, since fruit set was similar between these populations. The total population inaccuracy was 21.56 mm2, with inaccuracy of low organs being higher (12.03 mm²) than that of high organs (9.53 mm²). Floral traits may have different effects on the reproductive success of morphs of distylous species; the corolla length of P. nuda only affected the reproductive success of short-styled flowers. Large corollas showed greater reproductive success and lower individual inaccuracies for this morph, indicating compensation for the effects of imperfect reciprocal herkogamy on reproductive success.

The function of the reciprocal position of anthers and stigmas of the floral morphs in distylous flowers is to promote cross-pollination between morphs (legitimate pollination). Deviations from reciprocity can negatively affect the rates of legitimate pollen deposition, but other floral traits or the pollination environment might contribute to the function of distyly. In populations of four species of Palicourea and Psychotria, we quantified reciprocity using adaptive inaccuracy and measured the stigmatic lobe length to estimate how these traits influenced legitimate pollen deposition. We described the functional groups of pollinators and compared legitimate pollen flow between morphs in populations visited by different groups. Variations in reciprocity and stigmatic lobe length correlated with the rates of legitimate pollen depositions, with more reciprocal stigmas and longer stigmatic lobes presenting proportionally more legitimate pollen. Populations most frequently visited by bees or with generalized pollination displayed asymmetries in pollen flow between morphs, but with higher legitimate pollination in the S-morph. In contrast, L- and S-flowers showed similar legitimate pollen deposition in the population predominantly visited by hummingbirds. Our results suggest that reciprocity affects the function of distyly; however, floral traits such as stigmatic lobe length and the pollination environment can also influence the rates of legitimate pollen.

Species of Cyclopia are used to produce honeybush tea, a uniquely South African beverage that is rapidly growing in popularity locally and abroad. In response to warnings about ecological sustainab...

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 ch...

Studies of floral polymorphisms have focused on heterostyly, while stigma-height dimorphism has received considerably less attention. Few studies have examined the reproductive biology of species with stigma-height dimorphism to understand how factors influencing mate availability and pollen transfer are related to morph ratios in populations. Floral morphological traits, especially herkogamy and reciprocity, pollinator visitation, breeding system and spatiotemporal mate availability, are known to affect inter-morph pollination and morph ratios in species with stigma-height dimorphism. In this study, we investigated the presence of stigma-height dimorphism and estimated morph ratios in four naturally occurring populations of Jasminum malabaricum. We quantified morph- and population-specific differences in the abovementioned factors in these populations to understand the observed morph ratios. The positions of anthers and stigmas were characteristic of stigma-height dimorphism, the first report of this polymorphism in the genus. All study populations were isoplethic, implying equal fitness of both morphs. Herkogamy was higher in the short-styled morph, while reciprocity was higher between the long-styled stigma and short-styled anthers. Long- and short-tongued pollinators were common floral visitors, and we observed no differences between morphs in spatiotemporal mate availability or pollinator visitation. Neither morph exhibited self- or heteromorphic incompatibility. The short-styled stigma had lower reciprocity but likely receives sufficient inter-morph pollen from long-tongued pollinators, and also by avoiding self-pollination due to higher herkogamy. These results highlight the importance of sufficient effective pollinators and floral morphological features, particularly herkogamy, in maintaining isoplethy in species with stigma-height dimorphism.

The genus Prepusa Martius is endemic to montane regions of Brazil and comprises only six species, most of them endangered. Although studies on reproductive biology are necessary to increase knowledge of this genus and to implement conservation plans, to date, no such studies have been conducted. This study assessed the floral biology, breeding system, and pollinators of Prepusa hookeriana, an endangered species of high-altitude grasslands. We perform field observations and manual pollination experiments. Certain floral traits of this species appear to be associated with hummingbird pollination, although ornithophily is unusual in Gentianaceae. Even though cross-pollination is favoured by the presence of movement herkogamy and protandry, both prior and delayed autonomous self-pollination were observed. The only pollinator observed was the hummingbird Amazilia versicolor. Prepusa hookeriana did not show pollen limitation in the year of study. The absence of pollen limitation is related to the high reproductive success of pollination due to visits, not to autonomous self-pollination. However, we share the idea that the ability of autonomous selfing can provide a backup to cope with occasional pollination failures.

The family Cleomaceae is characterized by remarkable floral diversity and abundant nectar and pollen production. In this study, we evaluated the flower functional characteristics associated with the floral diversity of 10 Cleomaceae species, in addition to respectively seven and five accessions of Tarenaya hassleriana and T. longicarpa. The flowers were examined through five types of crosses and thus we determined the most efficient type of crossing based on the number of seeds. In addition, we evaluated pollen limitation, self-incompatibility, self-pollination and explored the possible effects on seed germination. The species can be grouped into three groups based on sexual systems: andromonoecious, hermaphroditism (chasmogamous and cleistogamous), and polygamomonoecious flowers. Gynandropsis gynandra, T. aculeata, T. diffusa, and T. microcarpa exhibited autogamy, whereas the other species did not. Overall, hand cross-pollination produced the highest seed yield, followed by open pollination. Self-pollination produced the lowest number of seeds, with no seed production noted in 16 accessions. Interestingly, the pollination type did not affect seed germination. Together, these results indicate the potential of the Cleomaceae family as a source of information regarding mechanisms involved in the determination of reproductive traits in plants.

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.”

ispartof: JOURNAL OF ECOLOGY vol:108 issue:1 pages:81-93 status: published

PREMISE In plants, populations and species vary widely along the continuum from outcrossing to selfing. Life-history traits and ecological circumstances influence among-species variation in selfing rates, but their general role in explaining intraspecific variation is unknown. Using a database of plant species, we test whether life-history traits, geographic range position, or abundance predict selfing rate variation among populations. METHODS We identified species where selfing rates were estimated in at least three populations at known locations. Two key life-history traits (generation time and growth form) were used to predict within-species selfing rate variation. Populations sampled within a species' native range were assessed for proximity to the nearest edge and abundance. Finally, we conducted linear and segmented regressions to determine functional relationships between selfing rate and geographic range position within species. RESULTS Selfing rates for woody species varied less than for herbs, which is explained by the lower average selfing rate of woody species. Relationships between selfing and peripherality or abundance significantly varied among species in their direction and magnitude. However, there was no general pattern of increased selfing toward range edges. A power analysis shows that tests of this hypothesis require studying many (i.e., 40+) populations. CONCLUSIONS Intraspecific variation in plant mating systems is often substantial yet remains difficult to explain. Beyond sampling more populations, future tests of biogeographic hypotheses will benefit from phylogeographic information concerning specific range edges, the study of traits influencing mating system (e.g., herkogamy), and measures of abundance at local scales (e.g., population density).

The factors that facilitate successful colonization of islands should be especially evident where the establishment filter is strongest. Colonizers of small, remote oceanic islands should be initially rare, extremely mate-limited and often without pollinators. Hence, plant communities on such islands should reflect an establishment history in which young ‘naturalized’ species are most likely to display self-compatibility and autonomous selfing, whereas ‘indigenous’ species may exhibit more diverse reproductive strategies. To test this prediction, we characterized breeding systems of 28 species on Pohnpei, in the Federated States of Micronesia, a group of remote Pacific islands that are considered a global biodiversity hotspot. Three families with both naturalized and indigenous species were selected—Fabaceae, Malvaceae and Melastomataceae. Measurements included field observations of dichogamy/herkogamy and floral attraction traits, pollen:ovule (P:O) ratios and experimental hand-pollinations for self-compatibility and pollen limitation. Phylogenetic generalized least squares analyses tested for trait correlations between naturalized and indigenous species. Flowers of all 28 species were bisexual, and pollinator attraction features were common. Pollen:ovule ratios ranged from 9 to 557 (median = 87), and all 11 hand-pollinated species were self-compatible. All species had >5 ovules and, Plants on remote, oceanic islands should reflect an establishment history in which recently ‘naturalized’ colonists can both self-pollinate without pollinators and self-fertilize, whereas older ‘indigenous’ species may evolve outcrossing mechanisms. In the first study of plant breeding systems on Pohnpei island, Micronesia, flowers of 28 species all exhibited overlapping male and female phases and short anther–stigma distances. Low pollen:ovule ratios of 9–557 suggested self-fertilization is common, as revealed by hand self-pollinations of 11 species. Since indigenous were no different than naturalized species, the data support a strong establishment filter and selection for reproductive assurance on oceanic islands.

Domestication is an evolutionary process with an impact on plant reproduction. Many domesticated plants are self-compatible (i.e., they lack mechanisms to reject their own pollen), but few domesticated plants are fully or partially self-incompatible. We used the husk tomato, Physalis philadelphica, as a study model to investigate changes in the reproductive strategy of an annual partially self-incompatible plant during the process of domestication. Wild and cultivated populations of this species coexist in close proximity. These different populations present a high level of morphological and genetic variation associated with different degrees of domestication. We hypothesized that artificial selection favors self-compatibility in cultivated plants through changes in their reproductive strategy and some reproductive parameters associated with domestication. To test this hypothesis, we characterized the floral morphology and some reproductive parameters of weedy plants (wild plants), landraces (semi-domesticated plant), and commercial plants (domesticated plants). We conducted an artificial crossing experiment, germinated the seeds, and recorded seedling growth. Commercial plants had the largest flowers and the highest number of ovules. Yet, they did not differ in other reproductive parameters (e.g., herkogamy, size of pollen grains, stigmatic area, and pollen:ovule ratio) from landraces and weedy plants. Physalis philadelphica produced fruits by autonomous autogamy in the artificial crossing experiment. These fruits were the smallest and lightest fruits at all degrees of domestication; however, fruit set of autonomous autogamy was higher in weedy plants. In addition, fruit production was higher when weedy plants donated pollen to commercial plants. Although seeds produced by autonomous autogamy of weedy plants had a low germination percentage, their cotyledons and the embryonic foliage leaves appeared earlier than in landraces and commercial plants. In conclusion, the domestication syndrome in this plant was manifested as increments in flower size and ovule production. Contrary to expectations, there was higher fruit production by autonomous autogamy in weedy plants than in cultivated plants. It seems that artificial selection in P. philadelphica favors self-incompatibility in cultivated plants. Nonetheless, spontaneous self-pollination seems to be advantageous in weedy populations because they produced viable seeds from which cotyledons and the embryonic foliage leaves emerged earlier than in cultivated plants.

SUMMARY We have studied eight species classified within Primula L. subgenus Sphondylia (Duby) Rupr. Two have monomorphic flowers and four have typically distylous flowers. P. floribunda Wall, includes both monomorphic and distylous races, while P. boveana Decres. has an incompletely discontinuous dimorphism for style length and pollen size, but is monomorphic for anther position. All the species have been tested and they are fully self-fertile. Evidence suggests that dimorphisms for style length (G/g), pollen size (P/p) and anther position (A/a) are each under independent genetic control. In the monomorphic P. verticillata, variations in stigma papilla length and style cell length appear to result from developmental correlates with style-length variation. Both the monomorphic species show some continuous variation for herkogamy. Several of the species studied show an incomplete development of the full distylous syndrome typical of most Primula species. If it is assumed that these represent evolutionary stages in the development of the full syndrome, and if it is assumed that diallelic incompatibility evolved after the evolution of distyly, then it would appear that the components of the distyly linkage group are likely to have evolved from a non-herkogamous ancestor in the following order: (i) short style mutant G; (ii) small pollen mutant p; (iii) low anther position mutant a.

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, plantDatura inoxiain 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 betweenrandFewere 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 differenceFe–Fis 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 ofDatura 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.

The pollination syndrome hypothesis usually does not successfully apply to the diversity of floral phenotypes or help predict the pollinators of most plant species. In Bromeliaceae, there is a wide range of floral visitors, making its species ideal to test for a correlation between nectar and floral traits with pollination syndrome. In this study, we analyzed the floral features, nectar production patterns, pollinators and floral visitors of Vriesea gigantea, and discussed its potential adaptive and ecological significance. We study three natural populations from the Atlantic Forest, Southern Brazil. The species presented protogyny and herkogamy, and its anthesis occurred at different periods among different populations. Vriesea gigantea has a relatively constant rate of nectar production during the day that continues overnight but at a reduced rate. Newly opened flowers already have around 80.0 μl of nectar. Although classified as chiropterophilous, based on flower morphology and pollinator observations, our results show that hummingbirds are effective pollinators in the studied populations of V. gigantea. Resumo A hipótese da síndrome da polinização geralmente não se refere exatamente à diversidade de fenótipos florais ou ajuda a predizer os polinizadores da maioria das espécies de plantas. Em Bromeliaceae, podemos encontrar uma ampla gama de visitantes florais, tornando suas espécies ideais para testar uma correlação entre características florais e do néctar com síndromes de polinização. Neste estudo, analisamos características florais, padrões de produção de néctar, polinizadores e visitantes florais em três populações naturais da Mata Atlântica de Vriesea gigantea, e discutimos seu potencial adaptativo e significado ecológico. A espécie apresentou protoginia e hercogamia, e sua antese ocorreu em diferentes períodos entre as populações. Vriesea gigantea tem uma taxa constante de produção de néctar durante o dia, com uma quantidade reduzida durante a noite. Flores recém-abertas possuem um considerável acúmulo de néctar. Embora a espécie tenha sido classificada como quiropterófila com base na morfologia das flores e observação de polinizadores, nossos resultados mostraram que os beija-flores são os polinizadores prováveis nas populações estudadas de V. gigantea.

Within the theoretical framework of the correlation pleiades, floral phenotypic integration has been proposed as a consequence of selection mediated by pollinators acting on floral characters. Here, we analyzed that assumption by studying the floral biology and pollination of the late-winter species Narcissus bulbocodium L. We found that the flowers of N. bulbocodium are pollinator-dependent (mainly on Bombus terrestris) in terms of achieving optimal levels of seed production (xenogamy mean seed-to-ovule ratio 64%). Flowers are phenotypically integrated, and only the inclusion of the stigma within the corona seems to have a positive and significant influence on the deposition of the pollen. It has been hypothesized that by including the stigma within the corolla, the flower has some control over the contact between stigma and pollinators that could lead to an “ordered herkogamy” as a way to promote outcross and avoid self-interference. Therefore, herkogamy was also studied, and while most previous studies have assessed the evolutionary significance of herkogamy by considering its relationship with outcrossing rates, we approach this phenomenon from a novel direction assessing the relationship between a proxy for herkogamy and the precision of the pollination process. Our results seem to support the existence of an optimal herkogamy distance that could maintain maximum levels of both pollen export and (cross) pollen capture. On the basis of the broad variability of herkogamy that we have found in N. bulbocodium and other data in the literature, we discuss the universality of the adaptive origin of herkogamy. Ministerio de Ciencia, Innovación y Universidades (España) | Ref. CGL2006-13847-CO2-02 Xunta de Galicia | Ref. CITACA-ED431E201 European Commission | Ref. LIFE18 GIE/IT/000755

Ceratostigma willmottianum (Plumbaginaceae) is a perennial herb native to China. Many species of Plumbaginaceae have been reported to exhibit heterostyly. Determining the functional breeding system of C. willmottianum can improve our understanding of the reproductive ecology of heterostylous plants. We investigated the floral traits and pollen and stigma characteristics in a natural population, and artificial pollination was carried out in an artificial population. It was found that C. willmottianum was distylous with short (S)- and long (L)-styled morphs, did not exhibit precise reciprocal herkogamy and was partially self-compatible but primarily outcrossing. In the artificial pollination experiments, the pollen tubes reached the base of the style under intermorph pollination, whereas they rarely penetrated the style under intramorph pollination and self-pollination. Both the L and S morphs exhibited a high seed set after intermorph pollination, whereas the seed set under intramorph pollination was lower. Therefore, C. willmottianum may spread the chance of receiving pollen between the two morphs by nonreciprocal heterostyly, which may be a unique mode of ecological adaptation in Plumbaginaceae. We believe that our discovery could provide new ideas regarding the origin and evolution of heterostyly.

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.

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 t...

Background and AimsTheory 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.MethodsWe 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 ResultsAs 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.ConclusionsGenetically 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.

Reproduction is key to determine the success of “winner” species. Plants that have an autogamous reproductive system to ensure their reproduction, but which maintain the possibility of cross-pollination, are the most flexible. Here, we aimed at understanding how those mechanisms occur in Sida cordifolia , an introduced pantropical weed. We recorded self-pollination mechanisms, monitored flower anthesis, and carried out a functional morphometric analysis. Based on those data, we established an experimental design of controlled pollination to measure the reproductive success generated by each self-pollination mechanism. We found two spontaneous self-pollination mechanisms: a prior one, during pre-anthesis, and a delayed one, at the moment of flower closure. However, partial dichogamy and herkogamy maintained during anthesis enabled cross-pollination. The floral attributes that maintain these mechanisms are fixed in the population. Alone, self-pollination mechanisms provide half of the fruit production, but together they are comparable to natural and cross-pollination with nearly total success. We show, for the first time, the interaction between prior and delayed self-pollination mechanisms and discuss how their combination enables autogamy but maintains the possibility of cross-pollination. The similar floral morphology of species of the pantropical genus Sida and the diversified self-pollination mechanisms in the family Malvaceae lead us to believe that this process may occur worldwide. Weedy species can be models for studies of floral biology, as they may have unique floral mechanisms that make them “winners” in the context of biotic homogenization.

The secondary presentation of pollen consists of a foral mechanism where the presentation of pollen occurs in other foral structures in addition to the anther, in order to increase the precision of the dispersion of the pollen by the vectors. This study aims to describe the temporal dynamics of secondary pollen presentation, and morphological and morphometric characteristics in stages of pre- anthesis and anthesis in genotypes of fve natural Heliconia psittacorum populations. For the study of foral morphometry traits of length of the fower, stamen, stigma and height of flament insertion in the petal in bud and fower were measured. The foral morphology of pre-anthesis buds and fowers in anthesis, the presence or absence of characteristics such as herkogamy, region of flament insertion in the petal, region of stylar hairs, and of secondary pollen deposition were evaluated. Treatments of controlled pollinations, self-pollination, geitonogamy, cross-pollination, natural pollination and growth to pollen tube were sampled. Floral herkogamy and pollen transfer to the adhered hairs in the stylar region were clearly observed during anthesis, constituting the frst record of occurrence of secondary pollen presentation in Heliconiaceae. Pollen tube growth was inhibited in the stigmatic, style and basal regions of the pistil. Natural fruiting produced little or no fruit. The positioning of the stamens above the stigma, pollen viability and stigma receptivity during anthesis of H. psittacorum fowers may favor self-pollination. The stylar hairs observed in all H. psittacorum populations’ help in the retention of pollen grains. The low fruiting rate in controlled and natural pollinations suggest that the main propagation form of H. psittacorum in the study areas is based on asexual reproduction.

Premise of research. Phenotypic traits that consistently mediate species’ responses to environmental variation (functional traits) provide a promising approach toward generalizing ecological and ev...

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.

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.

Background: Secondary pollen presentation (SPP) is a phenomenon where pollen is relocated from within the anthers to other floral structures for presentation.Aims: We report a new mechanism of androecial SPP on anther hairs, which is involved in increasing reproductive fitness in Brandisia hancei.Methods: The morphological change of anther hairs during floral ageing was recorded, and pollen grains were quantified after anther hair removal to clarify the roles played by hairs in pollen retention. Additionally, fruit set and seed set were assessed to investigate the efficiency of anther hairs to reproductive success.Results: Young flowers of B. hancei showed approach herkogamy, while autonomous selfing was achieved by changes in relative positions of the stigma and anthers during floral ageing. In B. hancei, anthers developed characteristic hairs of epidermal origin, and hairs originated along the furrow between the paired pollen sacs and were used for SPP after anther dehiscence. Anther hairs increased pollen retention by over 70% in later floral-age stages, and seed set decreased by about 30% after anther hairs had been removed.Conclusions: In the process of autonomous selfing, SPP on anther hairs enhances both the male and female functions of the flowers of B. hancei.

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. Fil: Roldán, Joanna Soledad. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina. Universidad Nacional de Córdoba; Argentina Fil: Ashworth, Lorena. Universidad Nacional de Córdoba; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina

The evolution of self-incompatibility seems to be responsible for a great part of the evolutionary success of flowering plants by preventing self-fertilization and, hence, the effects of inbreeding. The recognition of incompatibility mechanisms allows us to explore their emergency and functionality and is particularly interesting in hermaphrodite plants due to the sexual conflict they experience. Here, we investigated some pre- and post-pollination barriers for self-reproduction in Habranthus gracilifolius Herb. In a natural population, we performed hand pollinations to check mating and incompatibility systems, measured the distance between anthers and stigma within flowers, and observed flower visitors. Our results evidence that this species is self-incompatible and reproduces via outcrossing (90% of crossed flowers set fruits). Self-pollen grains germinate on the stigma surface and are stopped at the beginning of the style, typical of gametophytic self-incompatibility. The species presents approach herkogamy–stigma is positioned in average 2.78 mm above stamens, which may reduce pollen self-interference. It is visited by a variety of insects, many acting as thieves. Our indirect measures of pollinator performance suggest that flower visitation frequency alone is not enough to evaluate visitors’ effects on fruit set. Herkogamy and gametophytic self-incompatibility prevent self-pollination and self-fertilization, respectively, and together with pollinators appropriated size and correct work of blossoms ensure the reproductive success of this species.

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 .

Floral herkogamy, or the spatial separation of reproductive structures, plays an important role in the evolution of plant mating systems. One form of herkogamy is enantiostyly, or the alternate presentation of stigmas to the left and right of the floral axis. Theoretically, enantiostyly increases outcrossing rate by promoting reciprocal pollen deposition and export, but empirical evidence of this is lacking. Understanding how pollinators interact with enantiostylous flowers will illuminate the mechanisms by which enantiostyly encourages outcross pollination. We quantified style deployment within and among plants, of Allionia incarnata L. (Nyctaginaceae), a species with a new kind of enantiostyly we call “aggregate enantiostyly”. A. incarnata is the first species known to have “compound blossoms”, with different flowers each containing a left- or right-handed style. We quantify the autonomous selfing rate and describe floral visitor interactions with this unique floral structure. We found equal frequency of left- and right-handed blossoms within and among plants. Common floral visitors, Hemiargus ceraunus and Halictus sp., interacted with either anthers or stigmas, but not both, indicating effective outcrossing at the flower level. Outcrossing produced more fruits than selfing. The order in which floral visitors interacted with reproductive structures indicates that enantiostyly may increase pollen export. However, we did not find evidence that enantiostyly affected pollen deposition. Our study provides an intriguing first look at the reproductive ecology of a species with a new unique form of herkogamy, and provides evidence on the adaptive significance of enantiostyly.

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.

Prosopis L. is particularly rich and important in the Chaco region where forest vegetation currently persists as fragments. Its species are very important for the conservation and management of Chaco vegetation, and understanding their phenology and reproductive biology is essential. Here, we studied the phenology and pollination biology of P. rubriflora Hassl. in the Brazilian Chaco compared to other species of the genus. Differing from other Prosopis species, P. rubriflora is evergreen, and most phenophases are nonseasonal and bimodal (except fruiting), with a continuous pattern. Prosopis rubriflora is andromonoecious, but only weakly, since plants produce relatively few staminate flowers. Flowers showed three floral stylar phenotypes: short, medium and long style. Short-style flowers are functionally staminate because the stigma is nonreceptive. To compensate for the small size of P. rubriflora red brush flowers, attraction is compressed and transferred to the spike, which functions as a collective pollination unit. This compression may favour self- and intraplant pollination, which may reduce and/or prevent the occurrence of protogyny and partial, i.e. temporal, herkogamy in hermaphrodite flowers. Prosopis rubriflora has generalist pollination, but differs from the other Prosopis species because it is also pollinated by hummingbird, most likely due to its “ornithophilous” characteristics. This species is the first of the genus recorded as pollinated by hummingbirds. Although the exotic bee Apis mellifera L. presents high frequency of visit, this bee is not effective pollinator, due to its foraging style that does not favour cross-pollen flow. Native insects and hummingbird species are main pollinators because they present trapline foraging.

PREMISE OF THE STUDY The evolution of multiple floral traits often underlies the transition from outcrossing to selfing. Such traits can influence the ability to self, and the timing at which selfing occurs, which in turn affects the costs of selfing. Species that display variation in autonomous selfing provide an opportunity to dissect the phenotypic changes that contribute to variability in the mating system. METHODS In a common garden, we measured dichogamy and herkogamy in 24 populations of the protandrous mixed-mating herb Campanula americana, and related these to autonomous fruit set (autonomy). We then measured the timing of self-pollen deposition and fruit production in populations with high and low autonomy, and determined whether pollen germinability across floral development contributes to variation in autonomy. KEY RESULTS Populations that transitioned more rapidly to female phase displayed elevated autonomous selfing, but herkogamy was unassociated with autonomous selfing. Selfing occurred more rapidly in highly autonomous populations because of greater self-pollen deposition early in female phase. Pollen germinability in low-autonomy populations remained constant across floral development, but in high-autonomy populations it increased after floral anthesis and was highest near the onset of female phase. CONCLUSIONS Reduced dichogamy, elevated self-pollen deposition, and higher pollen germination late in male phase contribute to both earlier selfing and greater selfing. These traits vary among populations, likely reflecting past selection on the mating system. While delayed selfing bears fewer fitness costs, the evolution of earlier selfing may be favored if self-pollen availability decreases over floral development.

Heterostyly is a polymorphism in which populations comprise two (distyly) or three (tristyly) floral morphs with reciprocal positioning of the height of the anthers and stigmas (reciprocal herkogamy). Such reciprocal herkogamy permits precise pollen placement on pollinators’ bodies and pollination success by promoting disassortative pollen transfer between floral morphs. Here, we aimed to understand how the different components of reciprocity relate to and differ from each other in distylous flowers, and whether factors, such as genus, morph-ratio variation, and corolla tube length influence the imprecisions in the sexual organs within populations. We gathered literature and original morphometrics data comprising 98 Palicourea and Psychotria populations from 44 species, the two largest distylous Rubiaceae genera, which differ in some floral features and pollination systems (hummingbirds vs. insects, respectively). We estimated reciprocity using new inaccuracy indices and standardization techniques, which allowed comparisons among a wide array of populations. Our results showed that maladaptive bias (i.e. departure from optimum reciprocity) of low organs was higher than the other decomposed inaccuracies indices tested. In addition, inaccuracy of low organs was higher than those of high organs. We did not find any relation of either genus, morph-ratio variation, or corolla tube length on general inaccuracy measurements. Apparently, pollinator-mediated selection on floral traits, imposed by different pollinators groups, did not differ markedly among the studied distylous Rubiaceae. This is the first study on inaccuracy components for such an ample array of populations and provides insights on the relative importance of morphological traits that optimize the functionality of distyly. We highlight the applicability of using standardized reciprocity indices for cross populational investigations, which may both support previous insights and reveal unknown patterns in distylous plants.

Plant populations invading new environments might compromise their fitness contribution to the next generation, because of the lack of native specialist pollinators and/or potential mates. Thus, changes in plant mating system and traits linked to it are expected in populations colonising new environments where selection would favour selfing and floral traits that maximise reproductive output. To test this, we studied native (Mexico) and non-native (Spain) populations of the obligate sexual reproducing annual weed Datura stramonium. Flower size, herkogamy, total number of seeds per plant, number of visits by and type of pollinators, and inbreeding depression were assessed in native and non-native populations. Finally, we measured phenotypic selection on corolla size and herkogamy in each population. Flower size and herkogamy showed wide and similar variation in both ranges. However, the largest average flower size was found in one non-native population whereas the highest average positive herkogamy was detected in one native population. On average, flowers in the native range received more visits by pollinators. Hawkmoths were the main visitors in the native populations while only bees were observed visiting flowers in Spain's populations. Only in the native range was inbreeding depression detected. Selection to reduce herkogamy was found only in one native population. Absence of both inbreeding depression and selection on floral traits suggest a change in mating system of D. stramonium in a new range where generalist pollinators may be promoting high reproductive success. Selection against deleterious alleles might explain the reduction of inbreeding depression, promoting the evolution of selfing.

Floral monosymmetry appears to be a derived condition, arising independently many times, with a multiplicity of forms, and related to both, pollinator and plant diversification. It reduces interference between sexual functions in flowers and increases cross-pollination and allogamy. But since the description and assessment of the functioning of those floral features depend on fresh material and field observations, the number of groups thoroughly observed is limited. In the mostly Neotropical and actinomorphic Melastomataceae, monosymmetric flowers seem to be derived and the resulting herkogamy may constitute a mechanism to avoid anther–stigma interference and self-pollination. Here we studied Tococa guianensis, a Neotropical widespread shrub or treelet well known for its leaf–domatia ant interaction, but still lacking details on its breeding biology. We studied a population in Uberlândia, Central Brazil, observing floral biology features, diversity of visitors and breeding system. Flowers showed diurnal anthesis and a single-day life span. Pollen was the only floral resource offered, and poricidal anthers were clustered in the centre of the flower, while style was bent away from flower axis creating a transient herkogamy and flower monosymmetry. Style moves towards the floral axis during the day facilitating pollination. To date we do not know other Tococa species displaying such stylar features, which may be an intermediate form between the actinomorphic Miconieae and specialized zygomorphic Tococa. Contrasting with previous studies, the self-incompatibility found in T. guianensis indicates that transient herkogamy may function to avoid stigma clogging with self-pollen and allow flexible use of specialized buzz-pollinating bees.

Premise of the study Darwin proposed that the reciprocal arrangement of anthers and stigmas in heterostylous plants promotes cross-pollination through pollen segregation on pollinators' bodies. The floral tube in most heterostylous species constrains the feeding posture of pollinators determining the site of contact with sex organs located at different heights within a flower. Here, we evaluate Darwin's hypothesis in tristylous Lythrum salicaria, a species with a partially tubular corolla, and examine the extent to which the location of sex organs within a flower influence compatible and incompatible pollination. We predicted that the proficiency of cross-pollination would increase for more inserted sex organs due to the restrictions imposed by the floral tube on pollinator positioning. Methods We used experimental trimorphic and monomorphic arrays and emasculated flowers to quantify intermorph pollen transfer and capture among all sex-organ heights, and estimated the contribution of intraflower self-pollination, geitonogamous self-pollination, and intramorph outcross pollination to total intramorph pollination. Key results As predicted, disassortative pollination varied significantly with sex-organ height and was highest for short-level organs and lowest for long-level organs. In monomorphic arrays, most intramorph pollination resulted from outcross pollination followed by intraflower and geitonogamous self-pollination. Conclusions Our results provide evidence for the Darwinian hypothesis. Reciprocal herkogamy promoted varying degrees of disassortative pollination with the magnitude strongly influenced by sex-organ height within a flower.

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.

Premise The timing of self-fertilization has potentially important consequences for the trajectory of mating system evolution, the opportunity for outcrossing, and the maintenance of genetic variation in populations. For primarily selfing taxa, it remains poorly understood as to how floral variation influences the opportunity for outcrossing and whether those attributes vary among populations across geographic ranges. Methods We examined variation in floral traits (herkogamy, protandry, flower size, stigma stage at anthesis, timing of stigma receptivity) in seven populations of Clarkia xantiana ssp. parviflora, a primarily selfing taxon, spanning from the western to eastern margins of its distribution. We also performed experimental emasculations and pollinations (followed by stigma severing) to quantify the extent of opportunities for outcrossing across flower development. Results There was marked among-population variation in all floral traits, particularly between far eastern and western populations. Emasculation experiments showed that the eastern populations had minimal autonomous selfing, but western populations had high rates of selfing within 24 h after anthesis. Population variation in autofertility was significantly predicted by floral trait variation, especially protandry and petal size. Conclusions Greater protandry both extended the time over which outcrossing could potentially occur and reduced the probability of autonomous selfing, suggesting that there may be a tradeoff that results in fitness loss when pollinator visits are not common. The east-west pattern of differentiation in some floral traits parallels that of postglacial range expansion, suggesting that selection on the mating system may have been strong in the process of range expansion.

Herkogamy, the spatial separation of sex organs in hermaphroditic plants, has been proposed as a mechanism to reduce self-pollination and the associated processes of inbreeding and gamete wastage. Longitudinal herkogamy is the most frequent type, with two subtypes: approach herkogamy (anthers below the stigma), which is associated with diverse pollinator arrays, and reverse herkogamy (anthers above the stigma), associated with specialized, long-tongued pollinators. By using a herkogamy index that varied continuously from negative (reverse herkogamy) to positive (approach herkogamy) values, we studied the effect of continuous variation in herkogamy on pollinator attraction, selfing capability and plant fitness across three populations of Lonicera implexa differing in the relative abundance of long-tongued vs. short-tongued pollinators. Reverse herkogamy was significantly more frequent in the population where long-tongued pollinators were dominant than in the other two populations. Agreeing with this, the main floral visitors of L. implexa individuals with small and large herkogamy index were, respectively, long-tongued and short-tongued pollinators. Spontaneous selfing was low and increased with increasing herkogamy index (i.e. with approach herkogamy), although most of it occurred when there was close distance between anthers and stigma. Fruit production was unrelated to the herkogamy index in the population with long-tongued pollinators, but it increased with approach herkogamy (higher herkogamy index) in the other two populations. In contrast, seeds of individuals with reverse herkogamy (smaller herkogamy indices) germinated better. In this species, continuous variation in herkogamy might function as a reproductive strategy, as different morphotypes might be favoured by different pollinator assemblages., By using a herkogamy index that varied continuously from negative (reverse herkogamy) to positive (approach herkogamy) values, this study evaluates the effect of continuous variation in herkogamy on pollinator attraction, selfing capability and plant fitness in Lonicera implexa populations differing in the relative abundance of long-tongued vs. short-tongued pollinators. The results show that different morphotypes might be favoured by different pollinator assemblages, and that continuous variation in herkogamy in this plant species might function as a reproductive strategy, optimizing seed quality when long-tongued pollinators are abundant, and increasing fruit set when they are rare.

Herkogamy is an effective way to reduce sexual interference. However, the separation of stigma and anther potentially leads to a conflict because the pollen may be placed in a location on the pollinator different from the point of stigma contact, which can reduce pollination accuracy. Floral mechanisms aiming to resolve this conflict have seldom been explored. The floral biology of protandrous Ajuga decumbens was studied to uncover how the herkogamy dilemma can be resolved. Flower anthesis was divided into male, middle, female and wilting phases. The positions of stigma and stamen were dissimilar in different flower development stages. We measured the distance of the stamen and stigma to the lower corolla lip at different floral phases, which was the pollinators’ approaching way. The pollen viability, stigma receptivity, pollen removal and pollen deposition on stigma were investigated at different phases. During the male phase, the dehisced anthers were lower than the stigma, located at the pollinators’ approaching way, and dispersed most pollen with high viability. As the flower developed, the anthers moved upwards, making way for pollen deposition during the female phase. Meanwhile, the stigma becomes receptive by moving into the way and consequently was deposited with sufficient pollen. The position exchange of the stamen and stigma created a dynamic herkogamy at the floral phase with different sexual functions. This floral mechanism effectively avoided sexual interference and maintained pollination accuracy. In Ajuga, the movement herkogamy might be of adaptive significance in response to the changes in the pollination environment., Herkogamy may successfully avoid sexual interference but may also lead to reduction of pollination accuracy. The floral mechanism of protandrous Ajuga decumbens has perfectly resolved the herkogamy dilemma by putting the stamen and stigma in the same place but at times of different sexual functions. The movement of the stamen and stigma in opposite directions results in position exchange in the male and female floral phases. The interactions of dynamic herkogamy and dichogamy not only completely avoid sexual interferences, but also ideally maintain pollination accuracy. This novel floral mechanism highly enriches our understanding on the evolution of herkogamy.

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., In this study we investigated 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. 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 reciprocity between the sex organs of morphs.

Breeding system, sexual system, temporal variation in sex expression and herkogamy were evaluated in seven herbaceous-shrubby communities from the Gran Sabana Plateau, Venezuela. This analysis was conducted considering the life form, substrate type, succulence, carbon metabolism, nutritional relation, successional stage, pollination system specificity and endemism of plant species. Of the 348 plant species studied, 73.8 % were hermaphrodite, 16.9 % were monoecious and 9.2 % were dioecious. Plant sexual systems such as dichogamy and herkogamy were associated with life form, nutritional relations, carbon metabolism and pollination systems. Most species were adichogamous, followed by protandrous and protogynous. Protandry was high for perennial herbs, annual herbs and trees, and protogyny was most frequent in perennial herbs. Protandrous and protogynous species were frequently anemophilous. Herkogamy was higher than non-herkogamy. Herkogamy was higher for trees, shrubs and liana; higher in monophilous and lower in anemophilous species. Most of the hermaphrodites were herkogamous and adichogamous species. In contrast, monoecy were commonly perennial herb and dichogamous species and frequently associated with anemophily. Dioecious species were trees and shrubs and with polyphilous pollination. Dioecy was the most frequent sexual system for endemic species. Hermaphrodite species were similarly distributed across plant communities. Monoecy was slightly higher for savanna and fallow than the other communities, and dioecy was higher for shrublands and secondary bushland. Most plant species were non-agamospermous, non-spontaneous self-pollinated and xenogamous. Partially self-incompatible dominated, followed by self-incompatible, partially cross-incompatible and the lowest frequency corresponded to cross-incompatible species. All these results are discussed in the context of evolutionary and ecological trends., Breeding system, sexual system, dichogamy and herkogamy taking place in seven herbaceous-shrubby communities from the Gran Sabana Plateau, Venezuela, showed diverse mating strategies associated with life forms, pollination system specificity, successional stage, endemism, vegetation structure and community isolation. High levels of outbreeding strategies such as dioecy, monoecy, non-agamospermy, non-spontaneous self-pollination, xenogamy and partial self-incompatibility occurred mainly in woody species from shrublands and secondary bushland. Inbreeding strategies, non-herkogamy, spontaneous self-pollination and partial cross-incompatibility are primarily associated with herbaceous life forms from disturbed communities. These reproductive attributes are discussed in the ecological and evolutionary context.

Background and aims The serial homology of floral structures has made it difficult to assess the relative contributions of selection and constraint to floral integration. The interpretation of floral integration may also be clouded by the tacit, but largely untested, assumption that genetic and environmental perturbations affect trait correlations in similar ways. In this study, estimates of both the genetic and environmental correlations between components of the hawkmoth pollination syndrome are presented for chasmogamous flowers of Ruellia humilis , including two levels of control for serial homology. Methods A greenhouse population for quantitative genetic analysis was generated by a partial diallel cross between field-collected plants. An average of 634 chasmogamous flowers were measured for each of eight floral traits that contribute to the hawkmoth syndrome. Genetic correlations (across parents) and environmental correlations (across replicate flowers) were estimated by restricted maximum likelihood. Key results Stigma height, anther height and floral tube length were very tightly integrated in their responses to both genetic and environmental perturbations. The inclusion of floral disc width as a control for serial homology suggests this integration is an adaptive response to correlational selection imposed by pollinators. In contrast, integration of non-homologous traits was low. Furthermore, when comparisons between the dimensions of serially homologous structures were excluded, the genetic and environmental correlation matrices showed little congruence. Conclusions The results suggest that hawkmoths have imposed strong correlational selection on floral traits involved in the deposition and removal of pollen, and that this is a consequence of stabilizing selection on the relative positions of stigmas and anthers in the face of substantial flower size variation. Low integration of other floral traits, and conflicting patterns of genetic and environmental correlations among these traits, suggest weak or no correlational selection within the range of variability expressed within a population.

It is well known that animals can exert strong selective pressures on plant traits. However, studies on the evolutionary consequences of plant-animal interactions have mainly focused on understanding how these interactions shape trait means, while overlooking its potential direct effect on the variability among structures within a plant (e.g. flowers and fruits). The degree of within-plant variability can have strong fitness effects but few studies have evaluated its role as a potential target of selection. Here we reanalyzed data on Ipomoea wolcottiana stigma-anther distance to test alternate mechanisms driving selection on the mean as well as on intra-individual variance in two years. We found strong negative selection acting on intra-individual variation but not on mean stigma-anther distance suggesting independent direct selection on the latter. Our result suggests that intra-individual variance has the potential to be an important target of selection in nature and that ignoring it could lead to the wrong characterization of the selection regime. We highlight the need for future studies to consider patterns of selection on the mean as well as on intra-individual variance if we want to understand the full extent of plant-animal interactions as an evolutionary force in nature. This article is protected by copyright. All rights reserved.