Bee boys and fly girls: Do pollinators prefer male or female umbels in protandrous parsnip,Trachymene incisa (Apiaceae)?

Abstract Complete dichogamy occurs when temporal separation prevents any overlap in male and female function within and among flowers of one or more inflorescences. Although dichogamy may increase outcrossing and prevent inbreeding, it also results in the presentation of inflorescences with different floral resources. Pollinators may prefer one gender over the other based on these differences, which can reduce the transfer of pollen to conspecific stigmas and reduce floral resources for effective pollinators. We investigated whether the insect visitors of Trachymene incisa (Apiaceae), an Australian herb demonstrating complete protandry at the umbel level, show a preference for male or female umbels. The male phase umbels present pollen and nectar, whereas the female phase umbels offer nectar only. Therefore, we expect pollen‐collecting insects to favour male umbels, whereas insects that forage only for nectar will favour female umbels. In natural patches that exhibited a male umbel bias, insects showed a preference for male phase umbels at Agnes Banks in 2003 and at Myall Lakes in both 2003 and 2004. By contrast, insects showed no preference for umbel phases, visiting umbels at a similar frequency to which they occurred, at Agnes Banks in 2004 and at Tomago in both 2003 and 2004. This suggests spatial and temporal variation in insect preferences for umbel phases that differ in floral rewards. In experimental arrays where the umbel gender ratio was equal, there were no significant differences between male and female umbels in terms of insect visitation during a foraging trip and mean foraging time per visit. The differing patterns of preference may be due to a differential response by insects when the umbel ratios vary, where a male bias in umbel genders leads to a preference for male umbels, whereas an equal umbel gender ratio leads to equal visitation to male and female phase umbels.     

Functional analysis of synchronous dichogamy in flowering rush, Butomus umbellatus (Butomaceae)

Dichogamy is one of the most widespread floral mechanisms in flowering plants and is thought to have evolved to reduce interference between pollen import and export within flowers, especially self-pollination. Self-pollination between flowers may also be reduced if dichogamy is synchronous among flowers on an inflorescence. The analysis of dichogamy at both levels requires that the sexual phases of individual flowers be defined functionally in terms of pollen deposition and removal. We conducted morphological and functional analyses to investigate the degree of dichogamy within flowers and the synchronicity of dichogamy between flowers within inflorescences in an emergent, aquatic monocot, flowering rush (Butomus umbellatus). Based on daily observations of the development of marked flowers, data on the schedule of anther dehiscence within flowers, and repeat surveys of floral sex ratios in three populations, individual flowers appear to be strictly protandrous. On average, each flower spends ～1 d in each of male and female phases with an intervening 1-d neuter phase during which there is no available pollen in anthers and stigmas are not yet exposed to receive pollen. Morphological criteria used to delimit the beginning and end of each of these three sex phases were validated by quantifying the temporal schedule of pollen removal from anthers and pollen deposition on stigmas. Experimental pollinations showed that the morphological changes marking the end of female phase are hastened by pollen deposition. At the umbel level, synchronous development within sequential cohorts of flowers reduced overlap of male and female sexual phases between flowers. On average (±1 SE), 72 ± 3% of flowers completed their female phase while no other flowers on the same umbel were in male phase. Computer simulations of umbel development showed that this value is significantly higher than expected if the timing of flower development within umbels was random (30 ± 1%). Surveys of floral sex ratios in three populations revealed that 87% of umbels were either unisexual male or female at any given time. Pollinators usually visited more than one flower in sequence when foraging on umbels, suggesting that synchronous dichogamy may be an adaptation to avoid geitonogamy. The adaptiveness of both flower- and umbel-level dichogamy is also suggested because both traits are expressed to a lesser extent in obligately clonal, triploid populations, where flowers do not make seeds and hence floral adaptations are not maintained by natural selection.     

雄全同株植物簇花芹花期性别分配与开花式样

簇花芹(Soranthus meyeri)是古尔班通古特沙漠中常见的、具雄全同株性系统的伞形科多年生早春短命植物。该文对簇花芹花期性比(两性花数/总花数)与植株大小的关系及其开花式样进行了研究, 重点对花期大小依赖的性别资源分配进行了讨论。结果表明: 2006-2008年簇花芹群体水平的性比分别为0.69 ± 0.03、0.62 ± 0.03和0.69 ± 0.02, 彼此间无显著差异( p > 0.05), 表明其性比是相对稳定的, 可能受遗传因素的控制。雄花生物量与花粉量均比两性花的小, 说明产生雄花比产生两性花所需资源少。一级复伞形花序比二级复伞形花序具有较多的两性花, 说明前者易从植株上获得资源用于增加雌性适合度; 而后者产生较多的雄花以避免在雌性功能上资源投入的浪费, 增大花展示以吸引更多传粉者来增加花粉输出总量, 提高其整体适合度。植株水平的性比与地上营养器官的生物量间呈正相关关系, 说明较大个体对雌性功能的投资较大, 雌性繁殖成功受资源限制。复伞形花序内各伞形花序几乎同时向心开放, 且所有两性花及花序均为雄性先熟, 雌雄阶段完全分离, 但一级复伞形花序比二级复伞形花序早开放约5天, 彼此开花重叠期约为1天。这些特征对于一级复伞形花序进行异株异花授粉以及植株内不同级别花序间的同株异花传粉、避免雌雄功能间的干扰具有重要意义。     

Effects of variation in inflorescence size and floral rewards on the visitation rates of traplining pollinators ofAralia hispida

Summary In field experiments withAralia hispida inflorescences, the following variables were manipulated: number of umbels per inflorescence, number of flowers per umbel, and amounts of pollen and nectar per flower. Visitation rates by bumble bees, the principal pollinators, were then observed. In the reward-variation experiments, bees appeared to learn the positions of nectar-rich shoots, and visited them significantly more often than nectar-poor shoots. They did not respond to similar variation in pollen production. The nectar preferences developed slowly after the treatments were imposed, and bees continued to favor sites that had been occupied by nectar-rich shoots even after the treatments were discontinued. Visitation rate was approximately proportional to flower number, making it unlikely that increases in inflorescence size produced a disproportionate gain in male reproductive success (a necessary condition in certain models for the evolution of dioecy). For a fixed number of flowers per inflorescence, bees preferred inflorescences with more umbels. In pairwise choice tests of male-phase and female-phase umbels of various sizes, bees preferred male-phase umbels and larger umbels; the preference for male-phase umbels is stronger in bees that had previously fed on male-phase umbels.     

A REEXAMINATION OF THE POLLEN-DONATION HYPOTHESIS IN AN EXPERIMENTAL POPULATION OF ASCLEPIAS EXALTATA

The evolution of large floral displays in hermaphroditic flowering plants has been attributed to natural selection acting to enhance male, rather than female, reproductive success. Proponents of the “pollen-donation hypothesis” have assumed that maternal resources, rather than levels of effective pollination, limit fruit set. We investigated the pollen-donation hypothesis in an experimental population of poke milkweed, Asclepias exaltata, where effective pollination did not limit fruit set. Specifically, we examined the effects of flower number per plant, and flower number per umbel on male reproductive success (number of fruits sired) and female reproductive success (number of fruits matured). In 1990, a paternity analysis was performed on fruits collected from 53 plants whose inflorescences were not manipulated. Flower number per plant was significantly correlated with male success, but not with plant gender. Flower number per plant was also significantly correlated with female success, but umbel number and stem number per plant together explained more than half (58%) the variation in female success. The percentage of fruit set was not significantly correlated with flower number per plant. Plants with large floral displays did not disproportionately increase in male reproductive success, relative to female success, as predicted by the pollen-donation hypothesis. In 1991, the effect of flower number per umbel on male and female reproductive success was investigated. Flower number per umbel was manipulated on four umbels per plant by removing flowers to leave 6, 12, or 18 flowers in each umbel. Plants with the largest umbels effectively pollinated twice as many flowers on other plants, but produced only 1.35 times as many fruits as plants with 6 and 12 flowers per umbel. Relative maleness of plants with large umbels was nearly twice that of small and medium umbels. Although these observations are consistent with the pollen-donation hypothesis at the level of umbels, they are problematic, because much of the variation in flower number per umbel exists within, rather than among, plants in natural populations. Thus, plants consist of both reproductively male (large) and female (small) inflorescences, which act to increase total reproductive success. It is therefore inappropriate to explain the evolution of large floral displays in milkweeds solely in terms of potential male reproductive success.     

Protandry,synchronized flowering and sequential phenotypic unisexuality in neotropical Pentagonia macrophylla (Rubiaceae)

Summary The neotropical shrub Pentagonia macrophylla Benth. (Rubiaceae) has protandrous two-day flowers. Synchronous development among flowers on a single individual results in sequential phenotypic unisexuality: the entire plant alternates gender from day to day. Pistillate flowers produce more nectar than staminate flowers at comparable hours, but this difference does not result in different paterns of visitation to male and female flowers by hummingbird pollinators. Rare periods of bisexuality occur due to asynchronous floral development within or between inflorescences and are always followed by return to a synchronized pattern of alternation of gender. Reestablishment of synchrony usually involves a timeframe shift in the pattern of flowering (i.e., a plant produces staminate flowers on days when it previously would have been pistillate). It is suggested that timeframe shifts occur in response to inadequate pollination and serve to either desynchronize the plant from neighboring conspecifics or to temporarily allow self-pollination.     

Nectar production rates of Asclepias quadrifolia: causes and consequences of individual variation

The causes and reproductive consequences of individual variation in nectar production rates within a population of Asclepias quadrifolia were investigated. Two parameters were correlated with nectar production rate per flower: the root weight of the plant and the number of flowers in the inflorescence (umbel). Nectar production increased with increasing root weight but levelled off after a root size of about 3 g was reached. Nectar production decreased with increasing umbel size, but only for umbels that were greater than average size. A total of 57% of the variance in individual nectar production could be explained by these two variables with root weight accounting for 67% of the explained variance. Root weight is a good indicator of a plant's energetic status, indicating the importance of available energy in determining quantity of nectar produced. About 30% of the energy devoted to flowering is utilized in nectar production. Nectar production was significantly correlated with the male component of reproductive fitness, pollinaria removal, but not with the female component, pollinia insertion. Since pod production is limited by resource availability rather than the number of pollinia insertions, nectar production in A. quadrifolia is most closely associated with the maximization of the male function.     

Manipulation of bee behavior by inflorescence architecture and its consequences for plant mating

Angiosperms display flowers in many three-dimensional arrangements, but the functional significance of this diversity is largely unknown. We examined influences of inflorescence architecture on pollination and mating by quantifying the responses of bumblebees to three architectures and then using these observations as the basis of a model that simulated pollen dispersal. On artificial panicles, racemes, and umbels, each with 12 identical flowers, bees visited one more flower, on average, on umbels than on panicles (with racemes being intermediate). In contrast to this weak response, the consistency of foraging paths among flowers differed strongly among architectures (raceme > panicle > umbel). The simulation model revealed limited differences in self-pollination and pollen export among the three inflorescence designs when all flowers presented and received pollen, because mating differences depended on only the number of flowers visited. In contrast, in simulations of inflorescences on which pollen receipt and presentation were segregated so as to minimize interference among flowers, the consistency of movement paths governed mating. In this case, racemes self-pollinated much less than umbels (with panicles being intermediate), and racemes exported much more pollen than umbels and panicles. These effects have diverse consequences for the evolution of inflorescence architecture, flower design, and sexual segregation.     

Andromonoecy and developmental plasticity in Chaerophyllum bulbosum (Apiaceae–Apioideae)

Background and Aims

Andromonoecy, the presence of hermaphrodite and male flowers in the same individual, is genetically fixed or induced, e.g. by fruit set. Little is known about the forces triggering andromonoecy in the Apiaceae. In the present study, a natural population of the protandrous Chaerophyllum bulbosum was investigated to elucidate architectural constraints and effects of resource reallocation.

Methods

Three sets of plants (each n = 15) were treated by hand pollination, pollinator exclusion and removal of low-order inflorescences. Fifteen untreated plants were left as controls.

Key Results

Untreated plants produce umbels up to the third branch order, with increasing proportions of male flowers from 15 % (terminal umbel) to 100 % (third-order umbels). Fruit set correspondingly decreases from 70% (terminal umbel) to <10 % (second-order umbels). Insignificant differences from hand-pollinated plants do not reveal any sign of pollinator limitation at the study site. Bagged individuals show the same increase in male flowers with age as untreated plants, indicating that the presence of andromonoecy is not induced by fruit set. After umbel removal, individuals tend to present a higher number of hermaphrodite flowers and fruits in the umbels of second and third order. Three plants (25 %) produced an additional branch order composed of 100 % male umbels.

Conclusions

Inherited andromonoecy and the plastic response to environmental conditions are interpreted as a self-regulating system saving investment costs and optimizing fruit set at the same time.     

Bumblebee-pollination and temporal change of the calyx tube length in Clematis stans(Ranunculaceae)

Clematis stans is dioecious semi-arboreal, with pale purple–blue, nodding, tubulous flowers in a paniculate inflorescence. Both male and female flowers produce nectar from the base of the calyx tube during a flowering period of 3 or 4 days, and are pollinated by two bumblebee species, Bombus diversus and B. honshuensis, with different proboscis lengths. When the flowers open, four sepals constructing a calyx tube separate at the top and their respective tips gradually curl up, so that a tubular part shortens. Observations at two field sites showed that B. diversus (with a longer proboscis) most often visits the flowers with a longer calyx tube, and B. honshuensis (with a shorter proboscis) the flowers with a shorter calyx tube, i.e., later in the flowering period. By changing the calyx tube length, the flowers of C. stans accept the two bumblebee species with different proboscis length as pollinators and thus increase the chance of pollination for each flower. It was also found that the two bumblebee species prefer the male flowers to the female flowers, although the female flowers secrete more nectar as a reward than male flowers. This is likely because they visit the male flowers to collect pollen grains in addition to nectar. Electronic Publication     

新疆阿魏雄性不育的细胞形态学研究

新疆阿魏是特产于我国新疆的伞形科阿魏属多年生一次结实草本植物,属于国家二级保护的濒危植物。其种群中除了该科植物典型的雄全同株个体以外,还具有一定数量雄性不育的雌株。为了探究新疆阿魏的雄性不育现象及其影响因素,该文从细胞形态学角度对种群中的雌株以及雌花的形态特征进行了观测,采用石蜡切片技术对功能性雌花雄蕊的花药败育过程进行了观察。结果表明：(1)雌株3月底萌动,4月中旬进入花期,5月底果实成熟;物候期与雄全同株个体相同。(2)植株高度(71.00±10.92) cm和直径(71.67±17.64) cm、一级分支(23.83±2.04);基生叶长(33.41±11.63) cm、宽(24.47±8.60) cm;在植株大小、基生叶大小等方面雌株与雄全同株个体无差异。(3)在雌株上,一级分支和二级分支均为雌花序,均可结实;雌花序的伞幅数/每复伞花序(13.22±4.70)、花数/每花序(12.03±2.30)、总花数/每复伞花序(159.08)均高于两性花序;雌株比雄全同株个体产生更多可结实的花,形成更多具有杂种优势的异交后代。(4)雌花序中,花排列紧密,花间距小于两性花序和雄花序;开花时...     

Flowering, pollination, nectar standing crop, and nectaries ofEuterpe precatoria (Arecaceae), an Amazonian rain forest palm

Flowering and pollination biology of the monoecious palmEuterpe precatoria was studied in the forest of the Reserva Ducke near Manaus, Brazil, during two annual flowering periods (middle of October to end of March) between 1991 and 1993. Individuals produced one to four bisexual inflorescences per flowering period. Inflorescences have staminate and pistillate flowers in triads. The anthesis of the whole inflorescence averaged 26 days, with the male phase extending over the first 17 days, followed by six days without any open flowers, and the female phase covering the last three days. The distinct dichogamy of the inflorescence results in obligate outcrossing. The pollen-ovule ratio is high (42000). Both male and female flowers emit an almond oil-like scent and produce nectar in septal nectaries. The concentration and total amount of sugar of the nectar of female flowers (37% and 0.021 mg) were significantly higher than that of male flowers (9% and 0.007 mg). The nectar is hexose-rich and with a moderately high amino acid concentration (68 µg/ml).Euterpe precatoria seems to be a generally entomophilous palm with a predominance of beetles and bees as potential pollinators. The most constant visitors were beetles of the familiesCurculionidae, Chrysomelidae, Staphylinidae, and bees of the familyHalictidae. The actual composition of the insect spectrum depends on climatic and biotic factors. Additionally, wind pollination may occur. Fruit development lasted four months and ripe fruits are swallowed by toucans who thus disperse the seeds.This study is part of an unpublished doctoral thesis of the first author. The paper is dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer at the occasion of his 70th birthday     

Synchronized protandry and hermaphroditism in a tropical secondary forest tree, Schefflera heptaphylla (Araliaceae)

Selection favoring avoidance of stigma clogging, pollen discounting, self-fertilization, and other negative effects of self-pollination can produce intricate patterns of intra- and interfloral dichogamy in plants bearing numerous flowers. Here we report an extensive study of the relationships among dichogamy, floral sex allocation (pollen-to-ovule ratios), nectar production, floral visitors, mating system, and fruit set in natural populations of Schefflera heptaphylla, a widespread paleotropical secondary forest tree that produces thousands of flowers in a blooming season. Each tree produces 15?C30 sequentially blooming, paniculate, compound inflorescences. Each compound inflorescence has up to three orders of umbellets, which also bloom sequentially. While hand-pollinations showed that S. heptaphylla was capable of self-fertilization, our observations of thousands of flowers showed that strong intra- and interfloral protandry severely restricts both autogamous and geitonogamous self-pollination. All flowers were bisexual, thus the sexual system of the populations we studied was hermaphroditism. The pollen-to-ovule (P/O) ratios were characteristic of outcrossing species, and P/O ratios of flowers in the last-maturing (third order) umbellets were significantly higher than those in earlier-maturing (first and second order) umbellets. Floral visitors were primarily flies (Chrysomya sp. and Syrphinae sp.) and wasps (Vespula sp. and Eumenes sp.). Flowers produced nectar during both the male (pollen presentation) and female (stigma receptivity) stages of their development, and the volume of nectar production was higher in the female stage. Nevertheless, flowers received fewer visits in the female stage than they did in the male stage, and natural fruit set was low, especially in first and third order umbellets. Fruit set from hand cross- and self-pollinations was significantly higher than natural fruit set, indicating pollen limitation of fruit set. Schefflera heptaphylla has also been reported to be andromonoecious. Both hermaphroditism and andromonoecy are consistent with theoretical predictions for variation in sex allocation among sequentially maturing flowers in protandrous species. Further studies comparing hermaphroditic and andromonoecious populations of S. heptaphylla could elucidate the selective factors affecting sex expression, nectar production, and fruit set in species with numerous flowers displaying both intra- and interfloral dichogamy.     

Effect of nectar composition and nectar concentration on honey bee (Hymenoptera: Apidae) visitations to hybrid onion flowers

Floral nectar characteristics of nine inbred lines of onion (Allium cepa L.) were examined to determine their influence on the attractiveness of the onion flowers to honey bees (Apis mellifera L.). Potassium concentrations and sugar concentrations of the nectar did not significantly correlate with the number of bee visits received by an umbel. The average amount of nectar produced by both the umbels and the individual florets was significantly positively correlated with the number of bee visits. Our results suggest that selection for flowers with high nectar production may lead to a higher rate of pollination of the onion seed crop.     

Combined effects of inflorescence architecture, display size, plant density and empty flowers on bumble bee behaviour: experimental study with artificial inflorescences

Pollen dispersal by pollinators is governed by the extent to which diverse effects on pollinator behaviour act independently or augment or moderate each other. Using artificial inflorescences, we assessed the behavioural responses of bumble bees to inflorescence architecture (raceme, panicle, and umbel), inflorescence size (7 or 13 flowers), inter-inflorescence distance and the proportion of empty flowers per inflorescence. The advantage of large inflorescences in terms of attractiveness was larger for racemes and umbels than for panicles, whereas the effect of inter-inflorescence distance on the number of successive probes was smaller for racemes than for panicles and umbels. The number of flowers probed per visit increased almost proportionally with display size when fewer flowers were empty, whereas the number increased less when many flowers were empty. Our results suggest that display size and the spatial arrangement of flowers and nectar within inflorescences can contribute to efficient pollination by affecting pollinator behaviour interactively.     

Main sugar composition of floral nectar in three species groups of Scrophularia (Scrophulariaceae) with different principal pollinators

In some angiosperm groups, a parallelism between nectar traits and pollination syndromes has been demonstrated, whereas in others there is not such relationship and it has been explained as due to phylogenetic constraints. However, nectar trait information remains scarce for many plant groups. This paper focuses on three groups of Scrophularia species, with different flower sizes and principal pollinators, to find out whether nectar sugar composition is determined by pollinator type or reflects taxonomic affinities. Since the species we examined have protogynous flowers, and gender bias in nectar sugar composition has been noted in few plant groups, we also investigated whether sexual phase influenced Scrophularia nectar composition. The sugar composition was found to be similar in all species, having high‐sucrose nectar, except for the Macaronesian Scrophularia calliantha, which was the only species with balanced nectar; this last kind of nectar could be associated with the high interaction rates observed between S. calliantha and passerine birds. The nectar sugar composition (high in sucrose) was unrelated to the principal pollinator group, and could instead be considered a conservative taxonomic trait. No gender bias was observed between functionally female and male flowers for nectar volume or concentration. However, sexual phase significantly affected sucrose percentage in the largest‐flowered species, where the female phase flowers had higher sucrose percentages than the male phase flowers.     

Consequences of nectar robbing for the fitness of a threatened plant species

The effect of nectar robbing on plant fitness is poorly understood and restricted to a few plant species. Furthermore, the available studies generally evaluate the effects of nectar robbing on female fitness, disregarding the male component. Here we measured the effects of the nectar-robbing bumblebees on male (measured as pollen analogue flow distance) and female (measured as seed production) reproductive success in the insect-dependent Polygala vayredae, a narrow endemic species from the pre-Pyrenees (Spain). Intense nectar robbing by bumblebees significantly reduced the nectar available to legitimate pollinators in the studied population, and this reduction affected both male and female fitness. Significant differences were observed in fluorescent dye dispersion between robbed and non-robbed flowers within the population. Fluorescent dyes from non-robbed flowers were dispersed to larger distances and over a larger number of flowers when compared with robbed ones. Moreover, significant differences were observed in both fruit set and seed ovule ratios between the two groups, with non-robbed flowers presenting higher reproductive outcomes. However, no effect on seed weight was detected among treatments. The data obtained suggest that in this species, nectar robbing has important indirect and negative effects on plant fecundity, through both male and female functions, due to a modification in the foraging behaviour of legitimate visitors.     

Male-biased nectar production in a protandrous herb matches predictions of sexual selection theory in plants

Nectar production may disproportionately benefit male relative to female pollination success. In such cases, sexual selection is often suggested as the cause of asymmetric benefits, yet sexual selection in plants-particularly plants with hermaphroditic flowers-is infrequently tested empirically. Here, I used a protandrous herb with male-biased nectar production (Chrysothemis friedrichsthaliana, Gesneriaceae) to test predictions from sexual selection theory. During three flowering seasons, I measured nectar production, pollinator visits, and male and female fecundity following different numbers of cross-pollination events. In accordance with sexual selection predictions, (1) nectar production was greater during the male phase by at least 65%; (2) visits by the main pollinator (hummingbird Phaethornis striigularis) were limiting for part of the season, indicating that plants had to compete for pollinator visits; (3) pollinators spent 53% more time per visit and made 86% more visits to male- vs. female-phase flowers, suggesting that nectar increased male more than female pollination success; and (4) female fecundity was maximized by one visit, whereas male fecundity continued to increase with additional visits. Autonomous self-pollination further reduced visit requirements for maximum female seed set. These findings match specific sexual selection predictions: they link an observable male bias in a secondary sexual trait (nectar) to positive responses of mating participants (pollinators), resulting in more mating opportunities for mate-limited males, relative to apparently resource-limited females. This field-testing of theoretical predictions provides unique evidence that sexual selection helps maintain nectar production patterns in this and, quite likely, other hermaphroditic plant species.     

Pollination ecology ofOxalis violacea (Oxalidaceae) following a controlled grass fire

Vernal grass fires may encourage profuse flowering in clonal, colonies ofOxalis violacea. Long-styled colonies appear to be more floriferous than short-styled colonies and set a greater number of capsules. Individual flowers of both morphs live one or two days, change position on their respective pedicels and advertise nectar concealed at the base of the floral throat. AlthoughDiptera, Hymenoptera, andLepidoptera forage for nectar, bees (Andrenidae,Anthophoridae, Halictidae, andMegachilidae) probably make the only effective pollen transfers between the two morphs. Both male and female bees may transport pollen of both morphs and short-tongued bees (e.g.,Augochlorella spp.,Dialictus spp.) may be more common but as effective as pollinators as long-tongued bees (e.g.,Calliopsis andreniformis andHoplitis spp.). The conversion rate of flowers into capsules is only 13–17%. The spreading style in the short-styled morph is interpreted as an adaptation restricting insect-mediated, self-pollination but encouraging bee-stigma contact during nectar foraging.     

Nectar parasitism of Asclepias syriaca by ants: Effect on nectar levels,pollinia insertion,pollinaria removal and pod production

Summary Milkweed (Asclepias syriaca L.) umbels and stems attended by ants (Lasius neoniger Emery and Tapinoma sessile (Say)) initiated significantly fewer pods and showed a trend to produce fewer mature pods than did umbels and stems not attended by ants. Since similar numbers of pollinia were inserted in ant-attended flowers, we hypothesize that ant-excluded flowers obtain more pollinia from other clones than do ant-attended flowers of this normally non-selfing species. If pollinators from other clones first visited flowers not depleted of nectar by ants, they could provide this source of foreign pollinia. Our experiments suggest that nocturnal noctuid and geometrid moths can provide these pollinia.