MEASUREMENTS OF NATURAL SELECTION ON FLORAL TRAITS IN WILD RADISH (RAPHANUS RAPHANISTRUM). I. SELECTION THROUGH LIFETIME FEMALE FITNESS

Although the role of natural selection in the evolution of floral traits has been of great interest to biologists since Darwin, studies of selection on floral traits through differences in lifetime fitness have been rare. We measured selection acting on flower number, flower size, stigma exsertion, and ovule number per flower using field data on lifetime female fitness (seed production) in wild radish, Raphanus raphanistrum. The patterns of selection were reasonably consistent across three field seasons, with strong directional selection for increased flower production in all three years, weaker selection for increased ovule number per flower in two years, and selection for increased flower size in one year. The causes of the selection were investigated using path analysis combined with multiplicative fitness components. Increased flower production increased fruit production directly, and increased numbers of ovules per flower increased the number of seeds per fruit in all three years; pollinator visitation did not influence either of these fitness components. Increased flower size was associated with increases in both the number of fruit and the number of seeds per fruit in one year, with the latter relationship being stronger. Total lifetime seed production was affected more strongly by differences in fruit production than by differences in either the number of seeds per fruit or the proportion of fertilized seeds that were viable, but all three fitness components were positively correlated with total seed production.     

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.     

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.     

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.     

Environmental context drives seed predator-mediated selection on a floral display trait

Linking trait selection to environmental context is necessary to move beyond the simple recognition that selection is spatially variable and to understand what ultimately drives this variation. Natural selection acts through differences among individuals in lifetime fitness and information about effects on fitness components is therefore often not sufficient to gain such an understanding. We investigated how environmental context influenced intensity of seed predation, flower abortion and selection on floral display traits in 44–52 populations of the perennial herb Primula veris over 2 years. Phenotypic selection on both inflorescence height and flower number varied among populations and was mediated partly by pre-dispersal seed predation and flower abortion in one of the years. Among-population variation in selection on inflorescence height, but not flower number, was linked to variation in canopy cover via its effects on seed predation. Lifetime fitness was less sensitive to seed predator damage in shaded environments but estimates of selection based on lifetime fitness agreed qualitatively with those based on seed output. Our results demonstrate that seed predators constitute an important link between environmental conditions and trait evolution in plants, and that selection on plant traits by seed predators can depend on environmental context.     

Pollen limitation and natural selection on floral characters in the yellow monkeyflower, Mimulus guttatus

In flowering plants, pollen limitation has been proposed to intensify selection on floral characters important in pollinator attraction, but may also select for traits that increase seed set through autonomous selfing. Here, a factorial design (+/- pollen addition, +/- pollinator removal) was used to investigate how the pollination environment affects selection on floral morphology via female fitness in a mixed-mating population of the yellow monkeyflower, Mimulus guttatus (Phrymaceae). Female fitness was strongly pollen-limited, with supplementally pollinated plants setting 37% more seeds than open-pollinated individuals. Strong positive selection was found on flower length, weak positive selection on flower width : length ratio and no selection on stigma-anther distance in both open-pollinated and supplementally pollinated treatments. By contrast, flowers with relatively narrow corollas and low stigma-anther distances were favored in the pollinator exclusion treatment. These results provide mixed support for the idea that pollen limitation intensifies selection on floral characters. Despite strong phenotypic selection, natural pollen limitation did not mediate selection on characters associated with either pollinator attraction or self-fertilization. However, the novel pattern of selection on severely pollen-limited plants suggests that reproductive assurance against pollinator loss may have been directly involved in the floral evolution of closely related selfing taxa.     

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.     

Phenotypic selection in hawkmoth-pollinated Platanthera bifolia: targets and fitness surfaces

Abstract.— The present study explored phenotypic selection on phenological and morphological reproductive traits in hawkmoth-pollinated Platanthera bifolia (Orchidaceae), a Eurasian perennial herb displaying bisexual, long-spurred flowers. The work was carried out during three flowering seasons (1993–1995) in a Swedish population. Fitness was estimated as the number of pollinia removed (male fitness) and fruits produced (female fitness). Targets and patterns of selection were compared between years and sex functions by the use of multiple linear regression (including correlational selection estimates, i.e., of combination of traits), analysis of covariance, and projection pursuit regression (PPR). Results from the nonparametric surface-fitting-method PPR showed that selection was mostly linear, thus justifying the use of the parametric methods. In all study years, male and female fitness were highest in plants with many flowers. This reflects that flower number sets an upper limit to fitness and that a large inflorescence attracts more pollinators. In 1994, the summer was dry and the average spur length of P. bifolia was shorter than in the other years. In this year, male and female fitness were positively related to spur length, apparently because the spur of short-spurred plants was somewhat too short relative to the tongue length of the local pollinator for optimal pollen export and import. Additionally, the dry weather in 1994 caused a tendency for correlational selection, which was not found in the other years of study. Among small individuals (apparently more sensitive to drought than large ones), early-flowering plants had higher male and female fitness. The results show that patterns of selection may vary both between years and between sex functions in perennial hermaphroditic plants. The present study is one of the first to consider correlational selection in plants, which probably is of common occurrence and deserves to be investigated more.     

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.     

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.     

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。     

青藏高原高寒草甸植物花寿命与花吸引特征的关系及其对雌性繁殖成功的影响

花寿命指花保持开放且具有功能的时间长度, 是开花植物繁殖成功的一个重要功能性状。可塑性延长花寿命是植物在不利的传粉环境中保障繁殖的一种策略, 但延长花寿命也会增加繁殖成本。花寿命的可塑性变异不仅受传粉环境的影响, 而且还受资源分配权衡的影响。花寿命的理论模型指出, 植物的花寿命与花吸引特征之间存在资源分配权衡。为了研究在花粉限制环境中, 植物花寿命与花吸引特征之间的资源权衡及其对雌性适合度的相对重要性。该研究以青藏高原高寒草甸不同海拔(2 900和3 600 m)的11种开花植物为研究对象, 分析了不同植物群落中, 物种水平上: (1)花寿命与花吸引特征(花大小以及开花数目)之间的相关关系; (2)花寿命与花吸引特征对植物雌性适合度的相对贡献。结果表明, 无论是低海拔还是高海拔植物群落, 植物的花寿命与开花数目之间均存在权衡关系, 且长的花寿命增加了植物的雌性适合度。但在高海拔环境中, 植物的雌性适合度只与花寿命有关。这说明相对于低海拔植物, 花寿命对高海拔植物的雌性繁殖成功更为重要。     

Phenotypic selection in a natural population of Impatiens pallida Nutt. (Balsaminaceae)

In the annual plant Impatiens pallida, individuals exhibit a floral heteromorphism consisting of autogamously selfing, cleistogamous (CL) flowers and partially outcrossing, chasmogamous (CH) flowers. As part of an investigation into natural selection and mating system evolution in I. pallida, we measured the magnitude and direction of phenotypic selection on nine life history characters (two traits measured on three dates, one measured on two dates and one measured once). Three of these characters were positively correlated with the ratio of CH/CL flowers produced per plant, which is an important determinant of the mating system. Values for the nine characters and three different measures of fitness (viability, fecundity, lifetime) were estimated for 500 plants in five locations over a single growing season. Based on lifetime fitness, linear selection differentials were significant for all nine characters, indicating a selective advantage to tall, leafy, highly branched plants that flowered early. However, only two of these characters had a direct effect on fitness. Selection was significant on all nine characters when based on fecundity as well as lifetime fitness; however, only three of five characters examined had significant selection based on viability fitness. For all fitness components, the frequency of significant linear and nonlinear selection coefficients was comparable (23% vs 17% of all cases, respectively), but nonlinear coefficients were generally larger. Finally, the magnitude and direction of direct linear selection was heterogeneous among locations, for all characters and all fitness components. Collectively these results suggest that selection is strong, favouring large size, high allocation to reproduction and high CH/CL flower ratios. However, any directional evolutionary changes in vegetative or reproductive characters may be constrained by strong non-linear and correlational selection.     

Selection on floral traits through male fertility in a natural plant population

Most studies on selection in plants estimate female fitness components and neglect male mating success, although the latter might also be fundamental to understand adaptive evolution. Information from molecular genetic markers can be used to assess determinants of male mating success through parentage analyses. We estimated paternal selection gradients on floral traits in a large natural population of the herb Mimulus guttatus using a paternity probability model and maximum likelihood methods. This analysis revealed more significant selection gradients than a previous analysis based on regression of estimated male fertilities on floral traits. There were differences between results of univariate and multivariate analyses most likely due to the underlying covariance structure of the traits. Multivariate analysis, which corrects for the covariance structure of the traits, indicated that male mating success declined with distance from and depended on the direction to the mother plants. Moreover, there was directional selection for plants with fewer open flowers which have smaller corollas, a smaller anther–stigma separation, more red dots on the corolla and a larger fluctuating asymmetry therein. For most of these traits, however, there was also stabilizing selection indicating that there are intermediate optima for these traits. The large number of significant selection gradients in this study shows that even in relatively large natural populations where not all males can be sampled, it is possible to detect significant paternal selection gradients, and that such studies can give us valuable information required to better understand adaptive plant evolution. Second affiliation for Mark Van Kleunen is temporary until February 2007.     

GENETIC AND ENVIRONMENTAL VARIATION IN LIFE-HISTORY TRAITS OF A MONOCARPIC PERENNIAL: A DECADE-LONG FIELD EXPERIMENT

Directional and stabilizing selection tend to deplete additive genetic variance. On the other hand, genetic variance in traits related to fitness could be retained through polygenic mutation, spatially varying selection, genotype-environment interaction, or antagonistic pleiotropy. Most estimates of genetic variance in fitness-related traits have come from laboratory studies, with few estimates of heritability made under natural conditions, particularly for longer lived organisms. Here I estimated additive genetic variance in life-history characters of a monocarpic herb, Ipomopsis aggregata, that lives for up to a decade. Experimental crosses yielded 229 full-sibships nested within 32 paternal half-sibships. More than 5000 offspring were planted as seeds into natural field sites and were followed in most cases through their entire life cycle. Survival showed substantial additive genetic variance (genetic coefficient of variation ≈ 54%). Small differences at seedling emergence were magnified over time, such that the genetic variability in survival was only detectable by tracking the success of offspring for several years starting from seed. In contrast to survival, reproductive traits such as flower number, seeds per flower, and age at flowering showed little or no genetic variability. Despite relatively high levels of additive genetic variation for some life-history characters, high environmental variance in survival resulted in very low heritabilities (0–9%) for all of these characters. Maternal effects were evident in seed mass and remained strong throughout the lengthy vegetative period. No negative genetic correlations between major components of female fitness were detected. Mean corolla width for a paternal family was, however, negatively correlated with the finite rate of increase based on female fitness. That negative correlation could help to maintain additive genetic variance in the face of strong selection through male function for wide corollas.     

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.     

RESPONSES OF FLORAL TRAITS TO SELECTION ON PRIMARY SEXUAL INVESTMENT IN SPERGULARIA MARINA: THE BATTLE BETWEEN THE SEXES

Two widespread assumptions underlie theoretical models of the evolution of sex allocation in hermaphroditic species: (1) resource allocations to male and female function are heritable; and (2) there is an intrinsic, genetically based negative correlation between male and female reproductive function. These assumptions have not been adequately tested in wild species, although a few studies have detected either genetic variation in pollen and ovule production per flower or evidence of trade-offs between male and female investment at the whole plant level. It may also be argued, however, that in highly autogamous, perfect-flowered plant taxa that exhibit genetic variation in gamete production, strong stabilizing selection for an efficient pollen:ovule ratio should result in a positive correlation among genotypes with respect to mean ovule and mean pollen production per flower. Here we report the results of a three-generation artificial selection experiment conducted on a greenhouse population of the autogamous annual plant Spergularia marina. Starting with a base population of 1200 individuals, we conducted intense mass selection for two generations, creating four selected lines (high and low ovule production per flower; high and low anther production per flower) and a control line. By examining the direct and correlated responses of several floral traits to selection on gamete production per flower, we evaluated the expectations that primary sexual investment would exhibit heritable variation and that resource-sharing, variation in resource-garnering ability, or developmental constraints mold the genetic correlations expressed among floral organs. The observed direct and correlated responses to selection on male and female gamete production revealed significant heritabilities of both ovule and anther production per flower and a significant negative genetic correlation between them. When plants were selected for increased ovules per flower over two generations, ovule production increased and anther production declined relative to the control line. Among plants selected for decreased anthers per flower, we observed a decline in anther production and an increase in ovule production relative to the control line. In contrast, the lines selected for low ovules per flower and for high anthers per flower exhibited no evidence for significant genetic correlations between male and female primary investment. Correlated responses to selection also indicate a genetically based negative correlation between the production of normal versus developmentally abnormal anthers (staminoid organs); a positive correlation between the production of ovules versus staminoid organs; and a positive correlation between the production of anthers and petals. The negative relationship between male versus female primary investment supports classical sex allocation theory, although the asymmetrical correlated responses to selection indicate that this relationship is not always expressed.     

MEASUREMENTS OF SELECTION IN A HERMAPHRODITIC PLANT: VARIATION IN MALE AND FEMALE POLLINATION SUCCESS

I measured phenotypic selection of floral traits through both male and female functions of the hermaphroditic flowers of Ipomopsis aggregata (Pursh) V. Grant subsp. aggregata (Polemoniaceae). Fluorescent powdered dyes were used to track movement of pollen by hummingbirds and to measure pollen delivery to individual plants as well as pollen receipt. A phenotypic selection analysis revealed that selection due to male-male competition during pollination was capable of delaying flowering date and widening corolla tubes by 0.22 and 0.24 standard-deviation units, respectively, in a single generation. Several floral traits were highly correlated with each other. Multivariate selection analysis suggested that selection through male function directly favored late flowering as well as a sexual expression characterized by a short pistillate phase and long corollas. Selection intensities through male and female functions were of similar overall magnitude during the pollination stage of the life cycle, but different traits were favored, and selection sometimes acted in opposing directions. In 1985, selection through female function favored increased time spent in the pistillate phase and exserted stigmas (unlike selection through male function). As a result, individual plants varied greatly in functional gender. Plants that had exserted stigmas and narrow corollas and that spent a disproportionately long time in the pistillate phase achieved greater pollination success as females, while plants with the opposite traits achieved greater success as males. Moreover, female pollination success tended to increase, and male pollination success to decrease, with time spent in the pistillate phase, supporting a critical assumption of sex-allocation theory. Selection in the populations studied fluctuated from year to year and was highly sex-specific.     

No evidence for a role of pollinator discrimination in causing selection on flower size through female reproduction

The preference for certain floral phenotypes by flower visiting animals may fuel the evolution of floral traits because variation in flower visitation rates may lead to fitness variation within a population. Here, I examine the importance of flower size for pollinator visitation rate, seed set, and seed mass in two alpine populations of the insect-pollinated herb Ranunculus acris L. during two seasons. There was no pollen limitation of seed set or mass. Pollinators discriminated strongly against flowers experimentally reduced in size. Despite this, there were no signs of any significant impact of flower size on female reproductive success. The results show that although pollinators discriminate strongly among floral phenotypes, this may not always result in female fitness differences within a population because seed set or mass is not limited by pollen availability alone. Probably abiotic environmental constraints prevent plants with high pollinator visitation from capitalizing on the high pollen deposition.     

FERTILITY SELECTION ON A DISCRETE FLORAL POLYMORPHISM IN CLARKIA (ONAGRACEAE)

This study investigated fertility selection on a flower petal pigmentation polymorphism in Clarkia gracilis ssp. sonomensis. Natural populations are typically composed of nearly 100% spotted-petal plants, although rare populations contain a majority of unspotted plants. I compared fitness values for the two morphs using a simple fertility model to estimate selection for experimental arrays of plants placed into existing populations of different phenotypic frequencies. Both male and female reproductive success were estimated as well as the pattern of mating among phenotypes. Although the separate fitness components varied from no differences to a strong advantage for spotted plants, for every situation the selection calculations predicted an increase in the frequency of the spotted allele. Pollinator behavior and postpollination mechanisms may be responsible for the fitness differences. The apparent inability of the unspotted allele to spread though most natural populations is consistent with its selective disadvantage in this study.