Variation in resource limitation of plant reproduction influences natural selection on floral traits of <Emphasis Type="Italic">Asclepias syriaca</Emphasis>

The availability of both pollen and resources can influence natural selection on floral traits, but their relative importance in shaping floral evolution is unclear. We experimentally manipulated pollinator and resource (fertilizer and water) availability in the perennial wildflower Asclepias syriaca L. Nine floral traits, one male fitness component (number of pollinia removed), and two female fitness components (number of pollinia inserted and number of fruits initiated) were measured for plants in each of three treatments (unmanipulated control, decreased pollinator access, and resource supplementation). Although decreasing pollinators’ access to flowers did result in fewer pollinia inserted and removed, fruit set and phenotypic selection on floral traits via female and male fitness did not differ from the control. In contrast, resource supplementation increased fruit set, and phenotypic selection on seven out of nine floral traits was stronger via female than male fitness, consistent with the prediction that selection via female fitness would be greater when reproduction was less resource-limited. Our results support the hypothesis that abiotic resource availability can influence floral evolution by altering gender-specific selection.     

Effects of fire frequency on oak litter decomposition and nitrogen dynamics

Rapid speciation within some plant families has been attributed to the evolution of floral spurs and to the effect of spur length on plant reproductive success. The flowers of Impatiens capensis (jewelweed) possess a long, curved spur in which nectar is produced and stored. Spur length and curvature varies among plants within one population. Here I document that spur shape is variable in natural populations, variation within plants is less than variation among plants, and spur shape is correlated with components of female and male reproductive success. The apparent natural selection is weakly directional in 1 of 2 years, with greatest seed production and pollen removal occurring in flowers with the greatest spur curvature. Bee pollinator visit length is longest at flowers with highly curved spurs, and they leave less nectar in these spurs than in flowers with straighter spurs. Spur angle evolution may be limited, at least in part, by opposing selection by nectar-robbers who prefer to visit flowers with greater spur curvature. Other factors that might contribute to the maintenance of spur angle variation are temporal variation in the strength of selection and potential genetic correlations of spur shape with other traits under selection.     

The independent and combined effects of floral traits distinguishing two pollination ecotypes of a moth‐pollinated orchid

Identifying traits and agents of selection involved in local adaptation is important for understanding population divergence. In southern Sweden, the moth‐pollinated orchid Platanthera bifolia occurs as a woodland and a grassland ecotype that differ in dominating pollinators. The woodland ecotype is taller (expected to influence pollinator attraction) and produces flowers with longer spurs (expected to influence efficiency of pollen transfer) compared to the grassland ecotype. We examined whether plant height and spur length affect pollination and reproductive success in a woodland population, and whether effects are non‐additive, as expected for traits influencing two multiplicative components of pollen transfer. We reduced plant height and spur length to match trait values observed in the grassland ecotype and determined the effects on pollen removal, pollen receipt, and fruit production. In addition, to examine the effects of naturally occurring variation, we quantified pollinator‐mediated selection through pollen removal and seed production in the same population. Reductions of plant height and spur length decreased pollen removal, number of flowers receiving pollen, mean pollen receipt per pollinated flower, and fruit production per plant, but no significant interaction effect was detected. The selection analysis demonstrated pollinator‐mediated selection for taller plants via female fitness. However, there was no current selection mediated by pollinators on spur length, and pollen removal was not related to plant height or spur length. The results show that, although both traits are important for pollination success and female fitness in the woodland habitat, only plant height was sufficiently variable in the study population for current pollinator‐mediated selection to be detected. More generally, the results illustrate how a combination of experimental approaches can be used to identify both traits and agents of selection.     

Analysis of trait–performance–fitness relationships reveals pollinator-mediated selection on orchid pollination traits

Premise

The role of pollinators in evolutionary floral divergence has spurred substantial effort into measuring pollinator-mediated phenotypic selection and its variation in space and time. For such estimates, the fitness consequences of pollination processes must be separated from other factors affecting fitness.

Methods

We built a fitness function linking phenotypic traits of food-deceptive orchids to female reproductive success by including pollinator visitation and pollen deposition as intermediate performance components and used the fitness function to estimate the strength of pollinator-mediated selection through female reproductive success. We also quantified male performance as pollinarium removal and assessed similarity in trait effects on male and female performance.

Results

The proportion of plants visited at least once by an effective pollinator was moderate to high, ranging from 53.7% to 85.1%. Tall, many-flowered plants were often more likely to be visited and pollinated. Given effective pollination, pollen deposition onto stigmas tended to be more likely for taller plants. Pollen deposition further depended on traits affecting the physical fit of pollinators to flowers (flower size, spur length), though the exact relationships varied in time and space. Using the fitness function to assess pollinator-mediated selection through female reproductive success acting on multiple traits, we found that selection varied detectably among taxa after accounting for sampling uncertainty. Across taxa, selection on most traits was stronger on average and more variable when pollination was less reliable.

Conclusions

These results support pollination-related trait–performance–fitness relationships and thus pollinator-mediated selection on traits functionally involved in the pollination process.     

MEASUREMENTS OF NATURAL SELECTION ON FLORAL TRAITS IN WILD RADISH (RAPHANUS RAPHANISTRUM). II. SELECTION THROUGH LIFETIME MALE AND TOTAL FITNESS

It has often been suggested that selection on floral traits in hermaphroditic plants should occur primarily through differences in male fitness. However, measurements of selection on floral traits through differences in lifetime male fitness have been lacking. We measured selection on a variety of wild radish floral traits using lifetime male fitness measures derived from genetic paternity analysis. These male fitness estimates were then combined with estimates of lifetime female fitness of the same plants to produce measurements of selection based on lifetime total fitness. Contrary to the prediction above, there was no strong evidence for selection on floral morphology through male fitness differences in any of the three years of the study, but there was strong selection for increased flower size through female fitness differences in one year. The main determinant of both male and female fitness in all years was flower number; this lead to moderately positive correlations between male and female fitness in all three years.     

Selection on floral design in Polemonium brandegeei (Polemoniaceae): female and male fitness under hawkmoth pollination

Plant-pollinator interactions promote the evolution of floral traits that attract pollinators and facilitate efficient pollen transfer. The spatial separation of sex organs, herkogamy, is believed to limit sexual interference in hermaphrodite flowers. Reverse herkogamy (stigma recessed below anthers) and long, narrow corolla tubes are expected to promote efficiency in male function under hawkmoth pollination. We tested this prediction by measuring selection in six experimental arrays of Polemonium brandegeei, a species that displays continuous variation in herkogamy, resulting in a range of recessed to exserted stigmas. Under glasshouse conditions, we measured pollen removal and deposition, and estimated selection gradients (β) through female fitness (seeds set) and male fitness (siring success based on six polymorphic microsatellite loci). Siring success was higher in plants with more nectar sugar and narrow corolla tubes. However, selection through female function for reverse herkogamy was considerably stronger than was selection through male function. Hawkmoths were initially attracted to larger flowers, but overall preferred plants with reverse herkogamy. Greater pollen deposition and seed set also occurred in reverse herkogamous plants. Thus, reverse herkogamy may be maintained by hawkmoths through female rather than male function. Further, our results suggest that pollinator attraction may play a considerable role in enhancing female function.     

Within‐plant variation in reproductive investment: consequences for selection on flowering time

Variation among the leaves, flowers or fruit produced by a plant is often regarded as a nuisance to the experimenter and an impediment to selection. Here, we suggest that within‐plant variation can drive selection on other plant‐level traits. We examine within‐plant variation in floral sex allocation and in fruit set and predict that such variation generates variation in male success among plants, thereby driving selection on flowering time. We tested this prediction in a simulation model estimating selection on flowering time through male fitness when floral sex allocation and/or fruit set vary directionally among flowers on plants. We parameterized the model through a quantitative literature survey of within‐plant change in sex allocation. As predicted, within‐plant variation in floral sex allocation and in fruit set probability can generate selection on flowering time through male fitness. Declining fruit set from first to last flowers on plants, as occurs in many species, selected for early flowering onset through male fitness. This result was robust to self‐incompatibility and to varying returns on male versus female investment. Selection caused by declining fruit set was strong enough to reverse the selection for late flowering that can be caused by intrafloral protandry. Our model provides testable predictions regarding selection on flowering time through male fitness. The model also establishes the intriguing possibility that within‐plant variation may influence selection on other traits, regardless of whether that variation is under selection itself.     

Pollinators and nectar robbers cause directional selection for large spur circle in Impatiens oxyanthera (Balsaminaceae)

Nectar spurs have an important role in floral evolution and plant–pollinator coadaptation. The flowers of some species possess spurs curving into a circle. However, it is unclear whether spur circle diameter is under direct selection pressure from different sources, such as pollinators and nectar robbers. In this study, we quantified selection on some floral traits, such as spur circle diameter in Impatiens oxyanthera (Balsaminaceae) using phenotypic selection analysis and compared the relative importance of pollinators and nectar robbers as selective agents using mediation analysis. The study showed that pollinators caused significant selection on corolla length, spur curvature and spur circle diameter while nectar robbers only imposed strong selection on spur circle diameter. Pollinators favored flowers with large corolla, curly spurs and large spur circle while nectar robbers preferred flowers with small spur circle. More pollinator visits resulted in higher female reproductive success, while robbery reduced female fitness. Conflicting selection on spur traits from pollinators and nectar robbers was not found. Mediation analysis showed that selection on floral traits through nectar robbing was stronger than selection through pollination. The results suggested that pollinators and nectar robbers jointly mediated the directional selection for large spur circle, and nectar robbers caused stronger selection than pollinators on floral traits.     

展毛翠雀的花性状表型选择

传粉者的选择作用是花表型性状进化的重要驱动力, 解析选择作用的强度是理解花进化的关键。通过表型操控实验和表型选择研究能够分析花性状与其适合度的关系, 探究花性状的表型选择作用。为揭示花性状变化对雌性适合度的影响, 本研究处理展毛翠雀(Delphinium kamaonense var. glabrescens)花萼片大小, 并进行表型选择分析。结果表明: 人为减小展毛翠雀花萼片显著降低了传粉者的访花频率, 但是并没有影响种子产量(种子数和结籽率), 说明展毛翠雀花萼片的大小不影响种子产量, 可能主要吸引传粉昆虫输出花粉。通过雌性适合度(种子数量)估计表型选择梯度, 发现花萼片大小(长和宽)没有受到显著的直接选择梯度。但是, 花距长受到显著的线性选差和选择梯度, 表明花距的延长能够增加种子产量。本研究表明展毛翠雀花性状受到选择的作用, 但萼片和花距有不同的功能, 分别影响传粉者访问频率和种子产量。     

POLLINATOR-MEDIATED SELECTION ON FLORAL DISPLAY AND FLOWERING TIME IN THE PERENNIAL HERB ARABIDOPSIS LYRATA

The evolution of floral display and flowering time in animal-pollinated plants is commonly attributed to pollinator-mediated selection. Yet, the causes of selection on flowering phenology and traits contributing to floral display have rarely been tested experimentally in natural populations. We quantified phenotypic selection on morphological and phenological characters in the perennial, outcrossing herb Arabidopsis lyrata in two years using female reproductive success as a proxy of fitness. To determine whether selection on floral display and flowering phenology can be attributed to interactions with pollinators, selection was quantified both for open-pollinated controls and for plants receiving supplemental hand-pollination. We documented directional selection for many flowers, large petals, late start of flowering, and early end of flowering. Seed output was pollen-limited in both years and supplemental hand-pollination reduced the magnitude of selection on number of flowers, and reversed the direction of selection on end of flowering. The results demonstrate that interactions with pollinators may affect the strength of selection on floral display and the direction of selection on phenology of flowering in natural plant populations. They thus support the contention that pollinators can drive the evolution of both floral display and flowering time.     

Divergent selection on flowering phenology but not on floral morphology between two closely related orchids

Closely related species often differ in traits that influence reproductive success, suggesting that divergent selection on such traits contribute to the maintenance of species boundaries. Gymnadenia conopsea ss. and Gymnadenia densiflora are two closely related, perennial orchid species that differ in (a) floral traits important for pollination, including flowering phenology, floral display, and spur length, and (b) dominant pollinators. If plant–pollinator interactions contribute to the maintenance of trait differences between these two taxa, we expect current divergent selection on flowering phenology and floral morphology between the two species. We quantified phenotypic selection via female fitness in one year on flowering start, three floral display traits (plant height, number of flowers, and corolla size) and spur length, in six populations of G. conopsea s.s. and in four populations of G. densiflora. There was indication of divergent selection on flowering start in the expected direction, with selection for earlier flowering in two populations of the early‐flowering G. conopsea s.s. and for later flowering in one population of the late‐flowering G. densiflora. No divergent selection on floral morphology was detected, and there was no significant stabilizing selection on any trait in the two species. The results suggest ongoing adaptive differentiation of flowering phenology, strengthening this premating reproductive barrier between the two species. Synthesis: This study is among the first to test whether divergent selection on floral traits contribute to the maintenance of species differences between closely related plants. Phenological isolation confers a substantial potential for reproductive isolation, and divergent selection on flowering time can thus greatly influence reproductive isolation and adaptive differentiation.     

毛茛状金莲花不同花期的花特征和访花昆虫的变化及表型选择

为了研究植物生长季内开花时间对花特征表型选择的影响,我们以青藏高原东缘高寒草地的毛茛状金莲花Trollius ranunculoides)为实验材料,在生长季内不同开花时间(花前期、花末期)测定花特征,观察访花昆虫的类群和访花频率,生长季结束后收集种子.根据昆虫访花的喜好和季节内类群与访花频率的变化,分析了不同开花时间毛茛状金莲花的花特征与昆虫的选择;并用种子产量表示雌性适合度,估计了毛茛状金莲花的花特征在不同开花时间所受的表型选择.结果表明:不同花期植物的花特征有显著差异,相应的访花昆虫的类群和频率也存在差异,不同类群昆虫访花喜好也不一样.蜂喜好花瓣和花萼较宽、花茎短和花茎数少的个体,这正符合花前期的特征,因而蜂的访花频率在花前期较高;蝇对花特征没有明显的偏好.而通过雌性适合度估计毛茛状金莲花花特征所受的表型选择则是:花前期,花茎较长和花茎数多的植株适合度大;花末期,花茎数多的植株适合度大.我们的研究表明:在植物生长季,花期的分化伴随着传粉昆虫活动的变化.不同花期,访花昆虫的变化可能对植物花特征的分化起了至关重要的作用.但是访花昆虫对花特征的选择与通过雌性适合度估计植物受到的选择不尽相同,这可能是由于其他因素造成的.     

MULTIYEAR STUDY OF MULTIVARIATE LINEAR AND NONLINEAR PHENOTYPIC SELECTION ON FLORAL TRAITS OF HUMMINGBIRD-POLLINATED SILENE VIRGINICA

Pollination syndromes suggest that convergent evolution of floral traits and trait combinations reflects similar selection pressures. Accordingly, a pattern of selection on floral traits is expected to be consistent with increasing the attraction and pollen transfer of the important pollinator. We measured individual variation in six floral traits and yearly and lifetime total plant seed and fruit production of 758 plants across nine years of study in natural populations of Ruby-Throated Hummingbird-pollinated Silene virginica. The type, strength, and direction of selection gradients were observed by year, and for two cohorts selection was estimated through lifetime maternal fitness. Positive directional selection was detected on floral display height in all years of study and stigma exsertion in all years but one. Significant quadratic and correlational selection gradients were rare. However, a canonical analysis of the gamma matrix indicated nonlinear selection was common; if significant curvature was detected it was convex with one exception. Our analyses demonstrated selection favored trait combinations and the integration of floral features of attraction and pollen transfer efficiency that were consistent with the hummingbird pollination syndrome.     

Selection of trait combinations through bee and fly visitation to flowers of Polemonium foliosissimum

Pollinators are known to exert natural selection on floral traits, but the extent to which combinations of floral traits are subject to correlational selection (nonadditive effects of two traits on fitness) is not well understood. Over two years, we used phenotypic manipulations of plant traits to test for effects of flower colour, flower shape and their interaction on rates of pollinator visitation to Polemonium foliosissimum. We also tested for correlational selection based on weighting visitation by the amount of conspecific pollen delivered per visit by each category of insect visitor. Although bumblebees were the presumed pollinators, solitary bees and flies contributed substantially (42%) to pollination. In manipulations of one trait at a time, insects visited flowers presenting the natural colour and shape over flowers manipulated to present artificial mutants with either paler colour or a more open or more tubular flower. When both colour and shape were manipulated in combination, selection on both traits arose, with bumblebees responding mainly to colour and flies responding mainly to shape. Despite selection on both floral traits, in a year with many bumblebees, we saw no evidence for correlational selection of these traits. In a year when flies predominated, fly visitation showed a pattern of correlational selection, but not favouring the natural phenotype, and correlational selection was still not detected for expected pollen receipt. These results show that flower colour and shape are subject to pollinator‐mediated selection and that correlational selection can be generated based on pollinator visitation alone, but provide no evidence for correlational selection specifically for the current phenotype.     

Natural selection on floral traits through male and female function in wild populations of the heterostylous daffodil Narcissus triandrus

Measurements of natural selection in hermaphrodite populations require the analysis of performance through both female and male sex functions. Here, we investigate selection on three floral traits: flower number, flower length, and corona width through both sex functions in natural populations of the tristylous daffodil Narcissus triandrus . Selection through female function was examined in six populations, and in two of these we also estimated male selection gradients using multilocus microsatellite genotyping of parents and offspring. We detected significant directional selection for flower number through female function, and significant stabilizing selection for corona width and flower length through male function. Variation in male reproductive success was strongly influenced by the distance between mates and was significantly higher than variation in female reproductive success in one population, a result consistent with Bateman's principle. However, variation through both sex functions was similar in the other population and there was a significant negative correlation between female and male fitness indicating sex-specific trade-offs in reproductive success. Selection on floral design in N. triandrus was stronger through male than female function probably because floral morphology plays an important role in promoting effective cross-pollen transfer in populations of this heterostylous species.     

Altitudinal flower size variation correlates with local pollinator size in a bumblebee‐pollinated herb,Prunella vulgaris L. (Lamiaceae)

The influence of locally different species interactions on trait evolution is a focus of recent evolutionary studies. However, few studies have demonstrated that geographically different pollinator‐mediated selection influences geographic variation in floral traits, especially across a narrow geographic range. Here, we hypothesized that floral size variation in the Japanese herb Prunella vulgaris L. (Lamiaceae) is affected by geographically different pollinator sizes reflecting different pollinator assemblages. To evaluate this hypothesis, we posed two questions. (1) Is there a positive correlation between floral length and the proboscis length of pollinators (bumblebees) across altitude in a mountain range? (2) Does the flower–pollinator size match influence female and male plant fitness? We found geographic variation in the assemblage of pollinators of P. vulgaris along an altitudinal gradient, and, as a consequence, the mean pollinator proboscis length also changed altitudinally. The floral corolla length of P. vulgaris also varied along an altitudinal gradient, and this variation strongly correlated with the local pollinator size but did not correlate with altitude itself. Furthermore, we found that the size match between the floral corolla length and bee proboscis length affected female and male plant fitness and the optimal size match (associated with peak fitness) was similar for the female and male fitness. Collectively, these results suggest that pollinator‐mediated selection influences spatial variation in the size of P. vulgaris flowers at a fine spatial scale.     

蓝翠雀花退化雄蕊上的黄色髯毛对其繁殖成功的影响

蜜导是一类对传粉者具有特殊吸引和指示作用的花图案或花结构,在被子植物尤其是虫媒花中普遍存在,但目前有关蜜导对植物繁殖成功影响的研究相对较少.在野外条件下,我们以毛茛科翠雀属的蓝翠雀花(Delphinium caeruleum)为研究对象,通过人工授粉和去除髯毛实验对其交配系统和退化雄蕊上黄色髯毛的作用进行了初步研究,以探讨蜜导对于蓝翠雀花雌雄繁殖适合度的影响.结果发现,蓝翠雀花自交亲和,但其雌雄异熟和雌雄异位相结合的花特征完全避免了自动自交,因此该物种是专性异交植物,其传粉过程需要传粉者协助才能完成.与对照相比,去除髯毛处理显著降低了花粉的输出总量,但坐果率和结籽数并没有显著降低.我们的研究结果表明,蓝翠雀花退化雄蕊上的髯毛确实起到了蜜导的作用,但蜜导的存在主要提高了该植物的雄性繁殖适合度,表明雄性适合度的繁殖成功在蓝翠雀花的花进化过程中具有重要的作用,但这一结果仍有待在更多的居群中做进一步的验证.     

Variable selection in Platanthera bifolia (Orchidaceae): phenotypic selection differed between sex functions in a drought year

We estimated selection on three morphological characters in the hermaphroditic, hawkmoth-pollinated orchid Platanthera bifolia and explored selection surfaces through male and female function. The work was carried out in northern Sweden during two flowering seasons (1994 and 1995) in one natural population and one season (1995) in another natural population. Fitness was estimated as number of pollinia removed (male function) and number of fruits produced (female function). We detected directional selection towards larger inflorescence size (flower number) through both sex functions in both populations in 1995. In 1994, with an unusually dry growing season, 78% of the individuals failed to set any fruit, and there was selection for larger inflorescences only through male function. In this year, there was selection towards longer flower spurs, which could be a direct or indirect effect of spurs being shortened by drought. The results demonstrate that selection patterns may vary temporally and spatially, and that the 'male function hypothesis' may be applicable as female function is more resource dependent than male function.     

Manipulation of trait expression and pollination regime reveals the adaptive significance of spur length

Understanding the mechanisms of adaptive population differentiation requires that both the functional and adaptive significance of divergent traits are characterized in contrasting environments. Here, we (a) determined the effects of floral spur length on pollen removal and receipt using plants with artificial spurs representing the species-wide variation in length, and (b) quantified pollinator-mediated selection on spur length and three traits contributing to floral display in two populations each of the short-spurred and the long-spurred ecotype of the orchid Platanthera bifolia. Both pollen receipt and removal reached a maximum at 28–29 mm long spurs in a short-spurred population visited by short-tongued moths. In contrast, pollen receipt increased linearly across the tested range (4–52 mm) and pollen removal was unrelated to spur length in a long-spurred population predominantly visited by a long-tongued moth. The experimentally documented effects on pollen transfer were not reflected in pollinator-mediated selection through female fitness or pollen removal indicating that the natural within-population variation in spur length was insufficient to result in detectable variation in pollen limitation. Our study illustrates how combining trait manipulation with analysis of causes and strength of phenotypic selection can illuminate the functional and adaptive significance of trait expression when trait variation is limited.     

SELECTION ON POLEMONIUM BRANDEGEEI (POLEMONIACEAE) FLOWERS UNDER HUMMINGBIRD POLLINATION: IN OPPOSITION,PARALLEL, OR INDEPENDENT OF SELECTION BY HAWKMOTHS?

Particular floral phenotypes are often associated with specific groups of pollinators. However, flowering plants are often visited, and may be effectively pollinated by more than one type of animal. Therefore, a major outstanding question in floral biology asks: what is the nature of selection on floral traits when pollinators are diverse? This study examined how hummingbirds selected on the floral traits of Polemonium brandegeei, a species pollinated by both hummingbirds and hawkmoths. In array populations of P. brandegeei, we measured pollen movement, and female (seeds set) and male (seeds sired) fitness under hummingbird pollination. We then compared the patterns of selection by hummingbirds with our previous study examining selection by hawkmoths. We documented contrasting selection on sex organ positioning through female function, with hummingbirds selecting for stigmas exserted beyond the anthers and hawkmoths selecting for stigmas recessed below the anthers. Furthermore, hummingbirds selected for longer and wider corolla tubes, and hawkmoths selected for narrower corolla tubes. Therefore, contrasting selection by hawkmoths and hummingbirds may account for variation in sex organ arrangements and corolla dimensions in P. brandegeei. We documented how floral traits under selection by multiple pollinators can result in either an intermediate “compromise” between selective pressures (sex organs) or apparent specialization (corolla tube length) to one pollinator.