Mirror image flowers and their effect on outcrossing rate in Chamaecrista fasciculata (Leguminosae)

The influence of enantiostyly (reciprocal segregation of anthers and stigmas to different sides of the flower) on outcrossing rate was examined in Chamaecrista fasciculata (Leguminosae). I hypothesized that enantiostyly has not evolved to increase the female component of outcrossing and actually acts to increase the selling rate through geitonogamy. To quantify the role of enantiostyly to outcrossing, plants of known isozyme genotype were manipulated to be either completely left- or right-styled (nonenantiostylous) or to have equal numbers of left- and right-styled flowers (enantiostylous). Flower number was varied to quantify any interaction between floral display size and enantiostyly on outcrossing rate. These “target” plants were surrounded by unmanipulated plants homozygous for the alternative allele. Outcrossing rates of the target plants were determined by scoring the presence or absence of heterozygotes. The contribution of enantiostyly to geitonogamy may be reduced if pollinators discriminate among the floral types. Thus, observations of pollinator movement between flowers on the same plant were made to determine if pollinators discriminate between the floral types. Although pollinators moved randomly between flower types, outcrossing rate was only marginally effected by the presence of enantiostyly. Enantiostylous plants outcrossed at a slightly lower rate than nonenantiostylous plants only when the opportunity for geitonogamy was great. These results suggest that the contribution of enantiostyly to selling is minimal.     

The mechanism of delayed selfing in Collinsia verna (Scrophulariaceae)

Collinsia verna, blue-eyed Mary, has floral attributes of an outcrossing species, yet most flowers readily self-pollinate under greenhouse conditions. Here we describe the mechanism of self-pollination in C. verna via changes in relative positions of the stigma and anthers and late timing of receptivity, resulting in delayed selfing. Each flower contains four anthers that dehisce sequentially over ∼1 wk. Pollen that is not collected by pollinators accumulates in the keel petal and retains high viability (>80% pollen germination) up to the time of corolla abscission. The stigmatic surface does not become receptive until after the third anther dehisces. This overlap in the sexual phases is concurrent with a change in herkogamy during floral development. In most flowers (70%), the stigma has moved to the front of the keel and is positioned near the anthers when the third anther dehisces. Under field conditions, fruiting success of plants within pollinator exclosures was ∼75% of the fruiting success in open-pollinated plants (33% fruiting success via autogamy vs. 44% fruiting success, respectively). Collinsia verna plants in pollinator exclosures exhibit variation in autogamy rates within natural populations (range 0–80%). In addition, only half of naturally pollinated, receptive flowers examined had pollen tubes growing in their styles. In contrast, shortly after corolla abscission, nearly all flowers examined (96%) had pollen tubes in their styles. Thus we find that in C. verna, autogamy occurs late in floral development, which has the potential to provide substantial reproductive assurance, and that individuals vary in their ability to set fruit through this mechanism. We suggest that delayed selfing mechanisms may be overlooked in other species and that variable pollinator availability may play a significant role in the maintenance of mixed mating in species with delayed selfing, such as C. verna.     

FLORAL PARAMETERS,BREEDING SYSTEM AND POLLINATOR TYPE IN TRICHOSTEMA (LABIATAE)

Diversity in flower size and degree of exsertion of anthers and stigma from the corolla in the California species of Trichostema appear to be correlated with breeding system and pollinator type (bee vs. bird). Autogamous (self-pollinating) species unlike xenogamous (cross-pollinating) species lack spatially separate anthers and stigma and nototribic pollination. The outcrossing species have significantly larger flowers, significantly larger nectar volumes and significantly higher pollen-ovule ratios than do selfing species. Because autogamous species are less reliant on flower visitors to facilitate pollination, there may be relaxed selection for large nectar producing flowers. Pollen-ovule ratios are correlated with breeding system and reflect the efficiency of pollen transport. Data on floral parameters suggest xenogamous species expend more energy per floral unit and are less efficient seed producers than closely related autogamous species.     

Population-level variation in the expression of heterostyly in three species of Rubiaceae: does reciprocal placement of anthers and stigmas characterize heterostyly?

Heterostyly (i.e., reciprocal placement of anthers and stigmas between two or three floral morphs) is hypothesized to enhance outcrossing and reduce selfing. However, few studies have documented reciprocity among individual plants; instead, mean anther and stigma heights for floral morphs are usually reported, masking interindividual variation. We measured eight floral dimensions for individuals in five populations of three heterostylous Rubiaceae. The three methods used to quantify reciprocity yielded different conclusions regarding the degree to which populations conformed to expectations for heterostylous plants. Only Psychotria poeppigiana had stigma and, to a lesser degree, anther heights in discrete classes. Variation among plants of Bouvardia ternifolia and Psychotria chiapensis yielded a continuum of anther and stigma heights across populations. Comparison of distances between stigma and anthers indicated that only flowers of B. ternifolia had, as expected, a constant value for this distance. Finally, regression relationships between anther and stigma heights and corolla length showed that only in one population each of B. ternifolia and P. poeppigiana, and in P. chiapensis, was distance between anthers and stigmas the same across the range of corolla sizes for both floral morphs. Variation among these species in expression of heterostyly was not clearly linked to phylogenetic relationship or pollinator syndromes. Two approach herkogamous (AH) species were studied for comparison. Flowers of Psychotria brachiata were consistently AH, but flowers of P. pittieri were highly variable. Determining fitness consequences of population-level variation in sexual systems requires studies linking floral morphology to pollinator behavior and pollen transfer.     

山莨菪(茄科)的传粉生物学

茄科的多数种类具有自交不亲和的特点, 主要通过异花传粉结实; 但是, 一些物种或者物种内的部分种群或者个体却高度自交亲合, 转变为自交的繁育系统。该科植物山莨菪(Anisodus tanguticus)主要分布在青藏高原, 开花较早, 比其他晚开花的植物种类更加缺少有效的异花传粉昆虫。我们选择了位于不同海拔高度的2个种群进行比较研究, 主要目的是检验该物种的繁育系统是否在极端环境下由于传粉者的缺乏而发生了部分改变。研究发现,山莨菪的花不完全雌性先熟, 柱头和花药间的平均距离随着花开放时间的延长而不断缩小, 但两者在多数花的单花花期结束时并没有发生接触。因此, 山莨菪花主要表现为适应异花传粉的雌雄异位特征。然而, 少数花 (4.9％)的柱头和花药发生接触, 为“自动自交”的传粉解除了空间隔离。2个种群的多数个体存在自交不亲和机制, 应具有异花传粉的繁育系统; 但是部分个体具有明显的自交亲和能力, 为自交提供了生理基础。高海拔种群的传粉昆虫主要是厕蝇(Fannia sp.), 它们在不同植株间的活动能够保证异花传粉结实; 同时该种群的部分个体存在“自动自交”。低海拔种群的主要访花昆虫是蚂蚁, 它们在花内的活动导致花粉在同一朵花内传递, 而引起“协助自交”; 而异花传粉昆虫厕蝇的访花频率则较高海拔种群低。两个种群的结实均由于异花传粉者不足而受到传粉限制。因此两种不同类型的自交机制为该早期开花植物异花访花昆虫的不足提供了一定程度上的繁殖补偿。     

The effects of floral display on pollinator visitation vary among populations ofPhacelia linearis (Hydrophyllaceae)

Summary Individual plants in gynodioecious populations ofPhacelia linearis (Hydrophyllaceae) vary in flower gender, flower size, and flower number. This paper reports the effects of variation in floral display on the visitation behaviour of this species' pollinators (mainly pollen-collecting solitary bees) in several natural and three experimental plant populations, and discusses the results in terms of the consequences for plant fitness. The working hypotheses were: (1) that because female plants do not produce pollen, pollen-collecting insects would visit hermaphrodite plants at a higher rate than female plants and would visit more flowers per hermaphrodite than per female; and (2) that pollinator arrival rate would increase with flower size and flower number, the two main components of visual display. These hypotheses were generally supported, but the effects of floral display on pollinator visitation varied substantially among plant populations. Hermaphrodites received significantly higher rates of pollinator arrivals and significantly higher rates of visits to flowers than did females in all experimental populations. Flower size affected arrival rate and flower visit rate positively in natural populations and in two of the three experimental populations. The flower size effect was significant only among female plants in one experimental population, and only among hermaphrodites in another. The effect of flower number on arrival rate was positive and highly significant in natural populations and in all experimental populations. In two out of three experimental populations, insects visited significantly more flowers per hermaphrodite than per female and visited more flowers on many-flowered plants than on few-flowered plants, but neither effect was detected in the third experimental population. Because seed production is not pollen-limited in this species, variation in pollinator visitation behaviour should mainly affect the male reproductive success of hermaphrodite plants. These findings suggest that pollinator-mediated natural selection for floral display inP. linearis varies in space and time.     

Reproductive biology and pollinators in two enantiostylous Qualea species (Vochysiaceae) in the Brazilian Cerrado

• Enantiostyly is a floral polymorphism in which two floral forms in the same species differ in deflection of the stigma to right or left position. In monomorphic enantiostylous plants, flowers of the two morphs occur within the same individual, usually in the same proportion. In self‐compatible species the function of monomorphic enantiostyly is proposed to increase outcrossing rates and offer a reproductive advantage under pollination limitation. Enantiostylous species are usually self‐compatible and show heteranthery, with poricide anthers and pollen as pollinator reward; however, there are families, such as Vochysiaceae, that have different characteristics.
• We analysed the reproductive system and pollination biology of Qualea parviflora and Q. multiflora, two enantiostylous species from the Brazilian Cerrado that have specific morphological and physiological traits. For this, we characterized flower traits, performed hand pollinations and studied floral visitors.
• We found no differences between morphs in the proportion of flowers, nectar produced or its concentration, pollen quantity and fruit set. Both species were self‐incompatible and quite generalist regarding floral visitors.
• Enantiostyly in self‐incompatible plants seems to confer a reproductive advantage by reducing self‐interference resulting from stigma clogging. This novel result helps to expand our knowledge on this complex floral polymorphism and opens new avenues for future research on this topic.

     

Relative impact of mate versus pollinator availability on pollen limitation and outcrossing rates in a mass‐flowering species

Plant mating systems are driven by several pre‐pollination factors, including pollinator availability, mate availability and reproductive traits. We investigated the relative contributions of these factors to pollination and to realized outcrossing rates in the patchily distributed mass‐flowering shrub Rhododendron ferrugineum. We jointly monitored pollen limitation (comparing seed set from intact and pollen‐supplemented flowers), reproductive traits (herkogamy, flower size and autofertility) and mating patterns (progeny array analysis) in 28 natural patches varying in the level of pollinator availability (flower visitation rates) and of mate availability (patch floral display estimated as the total number of inflorescences per patch). Our results showed that patch floral display was the strongest determinant of pollination and of the realized outcrossing rates in this mass‐flowering species. We found an increase in pollen limitation and in outcrossing rates with increasing patch floral display. Reproductive traits were not significantly related to patch floral display, while autofertility was negatively correlated to outcrossing rates. These findings suggest that mate limitation, arising from high flower visitation rates in small plant patches, resulted in low pollen limitation and high selfing rates, while pollinator limitation, arising from low flower visitation rates in large plant patches, resulted in higher pollen limitation and outcrossing rates. Pollinator‐mediated selfing and geitonogamy likely alleviates pollen limitation in the case of reduced mate availability, while reduced pollinator availability (intraspecific competition for pollinator services) may result in the maintenance of high outcrossing rates despite reduced seed production.     

AN UNUSUAL KIND OF DISTYLY IN QUINCHAMALIUM CHILENSE (SANTALACEAE) ON VOLCÁN CASABLANCA,SOUTHERN CHILE

In mature flowers of the southern Andean parasitic herb, Quinchamalium chilense (Santalaceae), the stigmas and anthers are closer together in the pin morph than in the thrum morph. While the stigmas and anthers of the two morphs are in reciprocal positions as the flowers open, such reciprocity is lost as the result of post-anthesis allometric growth of the styles and stamens. Experimental pollinations reveal that both morphs of Q. chilense are self-compatible. Natural fruit set is higher on the pin morph. The latter is also statistically under-represented in natural populations with respect to a 1:1 ratio for pins and thrums. Pin flowers produce larger numbers of pollen grains than thrum flowers and pollen of pin flowers is smaller in size than that of thrum flowers. Higher fruit set on the pin morph is consistent with some tendency towards subdioecious breeding behavior, although an expected excess of geitonogamous pollinations on the pin morph might also be contributing to the difference in fruit set on the two morphs. The unusual floral morphology of Q. chilense could have evolved as a result of selection for larger flowers by certain pollinator groups less likely to effect geitonogamous pollination. Alternatively, extended development of post-anthesis flowers might be a reflection of selection for an array of flower sizes on individual inflorescences, producing a target effect for pollinator attraction. The evolution of such inflorescence morphology would have been facilitated by the fact that a trend in the direction of subdioecious breeding behavior accrues the same outcrossing advantages as strict distyly. Precise information on periods of stigma receptivity is required to further our understanding of the floral morphology of Q. chilense.     

Evolutionary consequences of ecological factors: pollinator reliability predicts mating‐system traits of a perennial plant

The reproductive‐assurance hypothesis predicts that mating‐system traits will evolve towards increased autonomous self‐pollination in plant populations experiencing unreliable pollinator service. We tested this long‐standing hypothesis by assessing geographic covariation among pollinator reliability, outcrossing rates, heterozygosity and relevant floral traits across populations of Dalechampia scandens in Costa Rica. Mean outcrossing rates ranged from 0.16 to 0.49 across four populations, and covaried with the average rates of pollen arrival on stigmas, a measure of pollinator reliability. Across populations, genetically based differences in herkogamy (anther–stigma distance) were associated with variation in stigmatic pollen loads, outcrossing rates and heterozygosity. These observations are consistent with the hypothesis that, when pollinators are unreliable, floral traits promoting autonomous selfing evolve as a mechanism of reproductive assurance. Extensive covariation between floral traits and mating system among closely related populations further suggests that floral traits influencing mating systems track variation in adaptive optima generated by variation in pollinator reliability.     

高山植物扁蕾的延迟自交机制

扁蕾(Gentianopsis barbata)具有鲜艳的花和显著的腺体,并且花开放的前5 d柱头和花药始终处于不同的位置(雌雄异位),这些花综合征表明该植物应为异花传粉。为检验这一假设,我们对青藏高原植物扁蕾的海北站种群进行了3年的传粉生物学研究实验。与花综合征所表明的繁育系统相反,两年的野外观察发现昆虫的访花频率十分低,不去雄并隔离昆虫处理也能产生大量种子,说明这一种群的繁殖主要是依赖于自花传粉。尽管利用种子结实评价的柱头可授性从花开放4 d后开始下降,但随着花的发育进程,雄蕊的伸长能使得花药与柱头完全接触。实验也证明,柱头可授性和花粉活力都超过5 d,说明花药和柱头的接触能够发生自花授粉。扁蕾的这种自花传粉机制应属于典型的延迟自交类型。自花授粉发生在单花花期快要结束前,自交之前仍然保持异交传粉机制,这种延迟自交避免了自交与异交竞争造成的花粉或者种子折损,并为扁蕾在青藏高原极端环境下由于访花昆虫缺乏造成的异交失败提供了繁殖保障。     

Impact of insect pollinator group and floral display size on outcrossing rate

Despite the strong influence of pollination ecology on the evolution of selfing, we have little information on how distinct groups of insect pollinators influence outcrossing rate. However, differences in behavior between pollinator groups could easily influence how each group affects outcrossing rate. We examined the influence of distinct insect pollinator groups on outcrossing rate in the rocky mountain columbine, Aquilegia coerulea. The impact of population size, plant density, size of floral display, and herkogamy (spatial separation between anthers and stigmas) on outcrossing rate was also considered as these variables were previously found to affect outcrossing rate in some plant species. We quantified correlations between all independent variables and used simple and two-factor regressions to determine direct and indirect impact of each independent variable on outcrossing rate. Outcrossing rate increased significantly with hawkmoth abundance but not with the abundance of any of the other groups of floral visitors, which included bumblebees, solitary bees, syrphid flies, and muscidae. Outcrossing rate was also significantly affected by floral display size and together, hawkmoth abundance and floral display size explained 87% of the variation in outcrossing rate. None of the other independent variables directly affected the outcrossing rate. This is the first report of a significant impact of pollinator type on outcrossing rate. Hawkmoths did not visit fewer flowers per plant relative to other pollinator groups but preferred visiting female-phase flowers first on a plant. Both the behavior of pollinators and floral display size affected outcrossing rate via their impact on the level of geitonogamous (among flower) selfing. Given that geitonogamous selfing is never advantageous, the variation in outcrossing rate and maintenance of mixed mating systems in populations of A. coerulea may not require an adaptive explanation.     

Does the morphological fit between flowers and pollinators affect pollen deposition? An experimental test in a buzz‐pollinated species with anther dimorphism

Some pollination systems, such as buzz‐pollination, are associated with floral morphologies that require a close physical interaction between floral sexual organs and insect visitors. In these systems, a pollinator's size relative to the flower may be an important feature determining whether the visitor touches both male and female sexual organs and thus transfers pollen between plants efficiently. To date, few studies have addressed whether in fact the “fit” between flower and pollinator influences pollen transfer, particularly among buzz‐pollinated species. Here we use Solanum rostratum, a buzz‐pollinated plant with dimorphic anthers and mirror‐image flowers, to investigate whether the morphological fit between the pollinator's body and floral morphology influences pollen deposition. We hypothesized that when the size of the pollinator matches the separation between the sexual organs in a flower, more pollen should be transferred to the stigma than when the visitor is either too small or too big relative to the flower. To test this hypothesis, we exposed flowers of S. rostratum with varying levels of separation between sexual organs, to bumblebees (Bombus terrestris) of different sizes. We recorded the number of visits received, pollen deposition, and fruit and seed production. We found higher pollen deposition when bees were the same size or bigger than the separation between anther and stigma within a flower. We found a similar, but not statistically significant pattern for fruit set. In contrast, seed set was more likely to occur when the size of the flower exceeded the size of the bee, suggesting that other postpollination processes may be important in translating pollen receipt to seed set. Our results suggest that the fit between flower and pollinator significantly influences pollen deposition in this buzz‐pollinated species. We speculate that in buzz‐pollinated species where floral morphology and pollinators interact closely, variation in the visitor's size may determine whether it acts mainly as a pollinator or as a pollen thief (i.e., removing pollen rewards but contributing little to pollen deposition and fertilization).     

FREQUENCY-DEPENDENT VARIATION FOR OUTCROSSING RATE AMONG FLOWER-COLOR MORPHS OF IPOMOEA PURPUREA

In a series of previous studies, it has been shown that populations of the common morning glory Ipomoea purpurea are typically polymorphic for flower-color genes that bias pollinator service and, hence, the rate of outcrossing. In this study, we show that the rate of outcrossing for the white flower-color morph depends on its frequency in experimental populations of the morning glory. White flowers are visited less often by the primary pollinator, bumblebees, and have lower outcrossing rates than colored flowers when they are in the minority. In contrast, blue or pink flower morphs are not undervisited and do not have lowered outcrossing rates when they are rare. In plant populations where genes that increase selling are selectively favored due to their transmission bias, undervisitation of rare morphs may act to stabilize genetic variation for outcrossing rates.     

BREEDING SYSTEM EVOLUTION IN LEAVENWORTHIA: BREEDING SYSTEM VARIATION AND REPRODUCTIVE SUCCESS IN NATURAL POPULATIONS OF LEAVENWORTHIA CRASSA (CRUCIFERAE)

Populations of Leavenworthia crassa (Cruciferae) studied for 3 years exhibited among- and within-population genetic variation for a suite of floral and reproductive traits (flower width, petal length, anther position, ability to set seeds in the absence of pollinators, time to first flowering) associated with breeding system. We used electrophoretic markers to show that a population with small, monomorphically colored flowers with introrse anthers had a significantly lower outcrossing rate (t = 0.03) than did a population with larger, polymorphically colored flowers with extrorse anthers (t = 0.33). In the more-outcrossing population the correlation between higher maternal plant outcrossing rate and the suite of six traits approached significance (P < 0.067), with greater petal size, greater flower width, and reduced ability to set seeds in the absence of pollinators contributing significantly. Plants in selfing populations had a generally higher reproductive success, with a higher number of flowers per plant, a smaller proportion of unfertilized ovules, a smaller proportion of fertilized ovules aborted, a higher rate of fruit set, and overall a larger number of seeds matured than did plants from the more outcrossing populations. Pollen limitation did not appear to account for lower reproductive success in outcrossing populations. Resource limitation did not differ substantially between populations. However, within-ovary patterns of fertilization, abortion, and seed weight were significantly less random in outcrossing populations than in selfing populations, suggesting that differential gamete and embryo success may be responsible for lower reproductive success in outcrossing populations.     

Modification of bumblebee behavior by floral color change and implications for pollen transfer in <Emphasis Type="Italic">Weigela middendorffiana</Emphasis>

Flowers of Weigela middendorffiana change the color from yellow to red. The previous study revealed that red-phase flowers no longer have sexual function and nectar, and bumblebees selectively visit yellow-phase flowers. The present study examined how retaining color-changed flowers can regulate the foraging behavior of bumblebees and pollen transport among flowers within (geitonogamous pollination) and between (outcrossing pollination) plants and how the behavior is influenced by display size (i.e., number of functional flowers) and visitation frequency. The visitation frequencies of bumblebees to plants and successive flower probes within plants were observed in the field using plants whose flower number and composition of the two color-phase flowers had been manipulated. To evaluate pollination efficiency over multiple pollinator visits, a pollen transport model was constructed based on the observed bumblebee behavior. In the simulation, three flowering patterns associated with display size and existence of color-changed flowers were postulated as follows: Type 1, large display (100 functional flowers) and no retention of color-changed flowers; Type 2, small display (50 functional flowers) and retention of color-changed flowers (50 old flowers), and; Type 3, large display (100 functional flowers) and retention of color-changed flowers (100 old flowers). Color-changed flowers did not contribute to increasing bumblebee attraction at a distance but reduced the number of successive flower probes within plants. Comparisons of pollen transfer between Types 1 and 3 revealed that the retention of color-changed flowers did not influence the total amount of pollen exported when pollinator visits were abundant (>100 visits) but decreased geitonogamous pollination. Comparisons between Types 2 and 3 revealed that the discouragement effect of floral color change on successive probes accelerated in plants with a large display size. Overall, the floral color change strategy contributed to reduce geitonogamous pollination, but its effectiveness was highly sensitive to display size and pollinator frequency.     

THE INTRAFLORAL PHENOLOGY OF STREPTANTHUS TORTUOSUS (BRASSICACEAE)

The sequence of floral events during anthesis was examined in Streptanthus tortuosus to determine the relationship between the male and female floral phases. The flowers are strongly protandrous. In the staminate phase, the anthers mature sequentially over a 3–4-day period. Because pollinators quickly remove pollen from the anthers, sequential anther maturation prolongs the male phase relative to what it would be if anthers did not mature sequentially. Pollen applied to the stigma during the staminate phase does not adhere readily and does not germinate. The length of the pistillate phase depends on pollinator activity, as pollination accelerates the abscission of floral parts. Unpollinated flowers remain pistillate for 3–4 days, during which time stigmatic receptivity declines gradually. In the field, 72% to 80% of flowers are staminate at any time, indicating that the staminate phase is three times longer than the pistillate phase when pollinators have access to the flowers. The consequences of the relative length of the floral phases and the schedule of stigmatic receptivity are discussed in terms of outcrossing mechanism, floral longevity, and sexual selection models.     

濒危植物长柄双花木的花部综合特征与繁育系统

 通过野外观察,运用杂交指数、花粉-胚珠比、人工授粉和套袋实验及等位酶电泳等方法,对长柄双花木(Disanthus cercidifolius Maxim. var. longipes H. T. Chang)自然种群和人工种群的开花状态、繁育系统进行了研究。结果如下：该种单花花期一般6～7 d,开花过程中柱头始终高于花药的位置,花药散粉具有先后顺序,一般2花药先散粉,其余3花药滞后1～2 d。单花花期依其形态和散粉时间可分为5个时期：即散粉前期、散粉初期、散粉盛期、凋谢期。种群开花一般历时49～55 d,但不同地点、年份之间有所差异,推测开花持续时间不同主要是环境条件所致。由于其杂交指数大于4,花粉-胚珠比等于1 250,结合人工套袋和授粉实验以及等位酶电泳结果可以确定该种的繁育系统属于异交为主,部分自交亲和,传粉过程需要传粉者。花粉竞争可能是导致该种濒危的主要的生殖生物学原因。     

Flower biology of the cactus Coryphantha elephantidens in the tropical dry forest of Central Mexico

Floral traits and sexual systems in angiosperms are strategies that enhance outcrossing within hermaphrodite flowers and among individuals in a population. Sexual systems with unisexual flowers have also evolved among angiosperms, resulting in sex specialization. Furthermore, the interaction of floral traits and floral visitors determines successful plant reproduction. Globose cacti are bee pollinated, and variation in the diversity of their pollinator assemblages is strongly associated with floral phenotype. Our objective was to describe the floral biology of the cactus Coryphantha elephantidens and to determine its relationship with pollinators. Floral traits were studied by direct observations in live and fixed flowers. The breeding system was determined using two estimators based on floral morphology: pollen grains to ovules per flower (P/O) ratio and outcrossing index. Pollination treatments were conducted to determine the mating system. Floral visitors were recorded using direct observation. Flowers of C. elephantidens are variable in color, protandric, herkogamous and nectarless. Estimators of the breeding system indicated xenogamy, which is consistent with the obligate outcrossing revealed by the pollination experiment. Thirty-seven percent of the plants have female flowers that do not produce pollen, making this population functionally gynodioecious. Both fruit and seed set were high compared to other globose cacti. Pollinators included eight species of native bees, a more diverse pollinator assemblage than other globose cacti. Given the high pressure on pollen due to functional gynodioecy, nectarless flowers, an outcrossing mating system, and the necessity of pollinators to set seeds, we concluded that native bees are highly efficient pollinators that play a crucial role in the sexual reproduction of C. elephantidens.     

Pollination ecology of Penstemon roseus (Plantaginaceae), an endemic perennial shifted toward hummingbird specialization?

Summary Pink-flowered tubular Penstemon roseus (Plantaginaceae), which has shifted partially to hummingbird pollination, blooms on high-elevation slopes in the mountains in Tlaxcala, Mexico. We studied the interactions between pollinator visitation rates to flowers, pollen removal and deposition, flower size, and nectar removal frequency on seed production in P. roseus. We combine observational and experimental studies in two contrasting natural populations. Our manual pollinations revealed that P. roseus is fully self-compatible. Autonomous self- and manual self-pollinated flowers matured as many seeds as when outcrossed, but outcrossing seems to become better than selfing as the flowering season progressed. Early in the season flowers that were bagged and hand-selfed, hand-outcrossed, or autonomously selfed, or unbagged and naturally pollinated had equal seed set in all four treatments. But later in the season, outcross pollen gave approximately twice as much seed set as the two self-treatments. Low levels of pollen receipt and pollen removal were consistent with the long time elapsed for a given plant to be visited by hummingbirds, which suggests pollen shortage in both sites. Despite differences in pollinator visitation rates to flowers, probability of flower visitation, removal and deposition of pollen, and nectar production rates between populations, we found that total nectar production had no effect on seed production at either site. The daily nectar secretion rate of 0.3–0.65 mg sugar per flower per 1–3 days was low relative to other hummingbird-adapted Penstemon species (typical range: 1.5–5 mg sugar per flower), and it might be intermediate between hummingbird- and bee-adapted Penstemon flowers. Our results support the hypothesis about a shift toward hummingbird pollination, and provide an example of a ‘despecialized’ Penstemon species, which attracts high-energy pollinators (hummingbirds) and profits from outcrossing, but retains bee-syndrome floral traits and low sugar production rates.