Effect of nectar supplementation on male and female components of pollination success in the deceptive orchid Dactylorhiza sambucina

Many orchids lack floral nectar rewards and therefore rely on deception to attract pollinators. To determine the effect that a mutation for nectar production would have on overall pollination success of the deceptive orchid Dactylorhiza sambucina, we recorded pollen deposition and removal in flowers of plants that had either been supplemented with an artificial nectar solution or left unmanipulated as controls. Nectar supplementation resulted in significant increases in the proportion of flowers pollinated, regardless of morph colour and the density of plants supplemented in the population. However, nectar supplementation had a significant positive effect on pollinaria removal only for the yellow morph in one experiment in which a low proportion of plants were supplemented. Thus a mutation for nectar production would have a positive effect on overall pollination success in D. sambucina, particularly the female component. The observed patterns are discussed in relation to other factors, such as cross-pollination and the reallocation of nectar resources for other plant functions, which are traditionally considered to shape the rewardless strategies of orchids.     

Pollination and breeding system of the enigmatic South African parasitic plant Mystropetalon thomii (Mystropetalaceae): rodents welcome,but not needed

• Unrelated plants adapted to particular pollinator types tend to exhibit convergent evolution in floral traits. However, inferences about likely pollinators from ‘pollination syndromes’ can be problematic due to trait overlap among some syndromes and unusual floral architecture in some lineages. An example is the rare South African parasitic plant Mystropetalon thomii (Mystropetalaceae), which has highly unusual brush‐like inflorescences that exhibit features of both bird and rodent pollination syndromes.
• We used camera traps to record flower visitors, quantified floral spectral reflectance and nectar and scent production, experimentally determined self‐compatibility and breeding system, and studied pollen dispersal using fluorescent dyes.
• The dark‐red inflorescences are usually monoecious, with female flowers maturing before male flowers, but some inflorescences are purely female (gynoecious). Inflorescences were visited intensively by several rodent species that carried large pollen loads, while visits by birds were extremely rare. Rodents prefer male‐ over female‐phase inflorescences, likely because of the male flowers’ higher nectar and scent production. The floral scent contains several compounds known to attract rodents. Despite the obvious pollen transfer by rodents, we found that flowers on both monoecious and gynoecious inflorescences readily set seed in the absence of rodents and even when all flower visitors are excluded.
• Our findings suggest that seed production occurs at least partially through apomixis and that M. thomii is not ecologically dependent on its rodent pollinators. Our study adds another species and family to the growing list of rodent‐pollinated plants, thus contributing to our understanding of the floral traits associated with pollination by non‐flying mammals.

     

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.     

Realized tolerance to nectar robbing: compensation to floral enemies in Ipomopsis aggregata

Background and Aims

Although the ecological and evolutionary consequences of foliar herbivory are well understood, how plants cope with floral damage is less well explored. Here the concept of tolerance, typically studied within the context of plant defence to foliar herbivores and pathogens, is extended to floral damage. Variation in tolerance to floral damage is examined, together with some of the mechanisms involved.

Methods

The study was conducted on Ipomopsis aggregata, which experiences floral damage and nectar removal by nectar-robbing bees. High levels of robbing can reduce seeds sired and produced by up to 50 %, an indirect effect mediated through pollinator avoidance of robbed plants. Using an experimental common garden with groups of I. aggregata, realized tolerance to robbing was measured. Realized tolerance included both genetic and environmental components of tolerance. It was hypothesized that both resource acquisition and storage traits, and traits involved in pollination would mitigate the negative effects of robbers.

Key Results

Groups of I. aggregata varied in their ability to tolerate nectar robbing. Realized tolerance was observed only through a component of male plant reproduction (pollen donation) and not through components of female plant reproduction. Some groups fully compensated for robbing while others under- or overcompensated. Evidence was found only for a pollination-related trait, flower production, associated with realized tolerance. Plants that produced more flowers and that had a higher inducibility of flower production following robbing were more able to compensate through male function.

Conclusions

Variation in realized tolerance to nectar robbing was found in I. aggregata, but only through an estimate of male reproduction, and traits associated with pollination may confer realized tolerance to robbing. By linking concepts and techniques from studies of plant–pollinator and plant–herbivore interactions, this work provides insight into the role of floral traits in pollinator attraction as well as plant defence.Key words: Compensation, herbivory, indirect effects, Ipomopsis aggregata, male reproductive success, nectar robbing, pollen donation, pollination, resistance, tolerance     

The influence of nectar production on hawkmoth behavior,self pollination,and seed production in Mirabilis multiflora (Nyctaginaceae)

Nectar is the most common floral reward that plants produce to attract pollinators. To determine the effect of nectar production on hawkmoth behavior, pollen movement, and reproductive success in Mirabilis multiflora, I manipulated nectar volumes and observed the subsequent foraging behavior of the hawkmoth Hyles lineata and the resulting pollen movement patterns. Individual hawkmoths visited significantly more flowers on plants with more nectar. The increase in flower visits significantly increased pollen deposition on stigmas and pollen removal from anthers when nectar volume was raised to twice the highest level found in nature. As hawkmoths visited flowers consecutively on a plant, the proportion of self pollen deposited on stigmas increased significantly and rapidly. Based on simulated hawkmoth visits, seed set was significantly reduced for flowers later in a visit sequence. A simple model combining these results predicts that the form of selection on nectar production varies depending on pollinator abundance. Using a multiple regression analysis a nearly significant (P < 0.08) effect of stabilizing selection was detected during a single season as predicted by the model for the prevailing hawkmoth abundance. Although increased nectar production may indirectly affect plant fitness by reducing resources available for other plant functions, the direct effect of high nectar production on pollinator behavior and self pollination may generally limit floral nectar production.     

The effects of nutrient addition on floral characters and pollination in two subalpine plants, <Emphasis Type="Italic">Ipomopsis aggregata</Emphasis> and <Emphasis Type="Italic">Linum lewisii</Emphasis>

The availability of soil and pollination resources are main determinants of fitness in many flowering plants, but the degree to which each is limiting and how they interact to affect plant fitness is unknown for many species. We performed resource (water and nutrients) and pollination (open and supplemental) treatments on two species of flowering plants, Ipomopsis aggregata and Linum lewisii, that differed in life-history, and we measured how resource addition affected floral characters, pollination, and reproduction (both male and female function). We separated the direct effects of resources versus indirect effects on female function via changes in pollination using a factorial experiment and path analysis. Resource addition affected I. aggregata and L. lewisii differently. Ipomopsis aggregata, a monocarp, responded to fertilization in the year of treatment application, increasing flower production, bloom duration, corolla width, nectar production, aboveground biomass, and pollen receipt relative to control plants. Fertilization also increased total seed production per plant, and hand-pollination increased seeds per fruit in I. aggregata, indicating some degree of pollen limitation of seed production. In contrast, fertilization had no effect on growth or reproductive output in the year of treatment on L. lewisii, a perennial, except that fertilization lengthened bloom duration. However, delayed effects of fertilization were seen in the year following treatment, with fertilized plants having greater aboveground biomass, seeds per fruit, and seeds per plant than control plants. In both species, there were no effects of resource addition on male function, and the direct effects of fertilization on female function were relatively stronger than the indirect effects via changes in pollination. Although we studied only two plant species, our results suggest that life-history traits may play an important role in determining the reproductive responses of plants to soil nutrient and pollen additions.     

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.     

Earlier snow melt and reduced summer precipitation alter floral traits important to pollination

Climate change can cause changes in expression of organismal traits that influence fitness. In flowering plants, floral traits can respond to drought, and that phenotypic plasticity has the potential to affect pollination and plant reproductive success. Global climate change is leading to earlier snow melt in snow-dominated ecosystems as well as affecting precipitation during the growing season, but the effects of snow melt timing on floral morphology and rewards remain unknown. We conducted crossed manipulations of spring snow melt timing (early vs. control) and summer monsoon precipitation (addition, control, and reduction) that mimicked recent natural variation, and examined plastic responses in floral traits of Ipomopsis aggregata over 3 years in the Rocky Mountains. We tested whether increased summer precipitation compensated for earlier snow melt, and if plasticity was associated with changes in soil moisture and/or leaf gas exchange. Lower summer precipitation decreased corolla length, style length, corolla width, sepal width, and nectar production, and increased nectar concentration. Earlier snow melt (taking into account natural and experimental variation) had the same effects on those traits and decreased inflorescence height. The effect of reduced summer precipitation was stronger in earlier snow melt years for corolla length and sepal width. Trait reductions were explained by drier soil during the flowering period, but this effect was only partially explained by how drier soils affected plant water stress, as measured by leaf gas exchange. We predicted the effects of plastic trait changes on pollinator visitation rates, pollination success, and seed production using prior studies on I. aggregata. The largest predicted effect of drier soil on relative fitness components via plasticity was a decrease in male fitness caused by reduced pollinator rewards (nectar production). Early snow melt and reduced precipitation are strong drivers of phenotypic plasticity, and both should be considered when predicting effects of climate change on plant traits in snow-dominated ecosystems.     

Spatially inconsistent direct and indirect effects of herbivory on floral traits and pollination success in a tropical shrub

Investigations on plant–animal interactions have traditionally focused on single interactions at a time (e.g. herbivory, pollination), yet plant fitness is generally influenced in complex ways by several interactions operating concurrently, and very little is known on the degree of spatial consistency of the direct and indirect effects that link different interactions. This paper evaluates experimentally whether direct and indirect effects of herbivory on male and female flower size and pollination success of the monoecious tropical shrub Cnidoscolus aconitifolius (Euphorbiaceae) remain consistent at three distant regions in Yucatan (southeastern Mexico). Plants were subjected to different levels of leaf defoliation, and treatment effects on floral traits (corolla area, corolla tube length, pollen production), and male and female components of pollination success (percent pollen removal, number of pollen tubes) were subsequently measured to evaluate the indirect effect of herbivory on plant reproductive success via pollination. Defoliation had significant direct effects on floral traits, but its indirect effects differed between the male and female components of pollination success. Moreover, the relationships between defoliation, floral traits and male and female pollination success varied spatially (i.e. between regions), although they were frequently more spatially‐consistent for male success than for female success. Results from this study stress the importance of explicitly testing for spatial variation in direct and indirect effects arising from plant‐animal interactions.     

INFLORESCENCE SIZE: TEST OF THE MALE FUNCTION HYPOTHESIS

One explanation for low fruit sets in plants with hermaphroditic flowers is that total flower production by a plant is controlled primarily by selection through male function. This male function hypothesis presupposes that success in pollen donation increases more strongly with flower number than does seed set. I tested this prediction by measuring male and female components of reproductive success as functions of flower number in natural populations of the self-incompatible, perfect flowered plant, Ipomopsis aggregata. Fruit set in this hummingbird-pollinated plant averaged 4.9 to 40.3% across the 4 years of study. Both the total amount of pollen donated and the total amount received, as estimated by movement of fluorescent powdered dyes, increased linearly with number of flowers on a plant. Total seed production, however, increased disproportionately quickly because plants with larger floral displays were more likely to set at least one fruit. An estimate of the functional femaleness of a plant, based on pollen donation and seed production, increased with flower number. These results do not support the male function hypothesis.     

Floral nectar production: what cost to a plant?

Floral nectar production is central to plant pollination, and hence to human wellbeing. As floral nectar is essentially a solution in water of various sugars, it is likely a valuable plant resource, especially in terms of energy, with plants experiencing costs/trade-offs associated with its production or absorption and adopting mechanisms to regulate nectar in flowers. Possible costs of nectar production may also influence the evolution of nectar volume, concentration and composition, of pollination syndromes involving floral nectar, and the production of some crops. There has been frequent agreement that costs of floral nectar production are significant, but relevant evidence is scant and difficult to interpret. Convincing direct evidence comes from experimental studies that relate either enhanced nectar sugar production (through repeated nectar removal) to reduced ability to produce seeds, or increased sugar availability (through absorption of additional artificial nectar) to increased seed production. Proportions of available photosynthate allocated by plants to nectar production may also indicate nectar cost. However, such studies are rare, some do not include treatments of all (or almost all) flowers per plant, and all lack quantitative cost–benefit comparisons for nectar production. Additional circumstantial evidence of nectar cost is difficult to interpret and largely equivocal. Future research should repeat direct experimental approaches that relate reduced or enhanced nectar sugar availability for a plant with consequent ability to produce seeds. To avoid confounding effects of inter-flower resource transfer, each plant should experience a single treatment, with treatment of all or almost all flowers per plant. Resource allocation by plants, pathways used for resource transfer, and the locations of resource sources and sinks should also be investigated. Future research should also consider extension of nectar cost into other areas of biology. For example, evolutionary models of nectar production are rare but should be possible if plant fitness gains and costs associated with nectar production are expressed in the same currency, such as energy. It should then be possible to understand observed nectar production for different plant species and pollination syndromes involving floral nectar. In addition, potential economic benefits should be possible to assess if relationships between nectar production and crop value are evaluated.     

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.     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

Sexual selection in a hermaphroditic plant through female reproductive success

Sexual selection is well accepted as a mechanism of shaping traits in animals. However, whether and how floral traits are sexually selected in hermaphroditic plants remains less clear. Here, we use Passiflora incarnata to address how floral traits that affect pollination success are selected via female function. We manipulated the ecological context by limiting pollination and adding resources to expand the phenotypic distribution and alter the intensity of sexual selection. Total sexual selection favoured lower style deflexion because of its impact on pollen receipt and subsequent seed number. However, total selection on style deflexion was not significant, indicating additional selection on style deflexion through routes other than mating. Limited pollination and enhanced resources were expected to alter the distribution of pollen deposition and seed production and therefore intensify the Bateman gradient – the relationship between pollen receipt and seed production. Indeed, the Bateman gradient was strongest when pollination was limited, suggesting potential for sexual selection to influence floral trait evolution under these conditions. Overall, we found floral traits may be shaped by sexual selection through female reproductive success in this hermaphroditic plant. These results support manipulations to enhance the variance in mating as a mechanism to understand patterns of sexual selection.     

Are nectar guide colour changes a reliable signal to pollinators that enhances reproductive success?

Background: Ageing and post-pollination changes in floral colour occur widely in flowering plants, but it remains an open question as to whether or not colour changes in nectar guides are associated with the quantity of floral rewards that ultimately influence pollinator visitations and reproductive success.

Aims: To examine whether nectar guide changes should be considered as a reliable signal to pollinators and to assess the effects of nectar guide changes on reproductive success.

Methods: We studied the process and adaptive value of colour changes in the nectar guides of Arnebia szechenyi whose flowers typically display conspicuous nectar guides at the onset of anthesis, after which they begin to fade, and disappear completely on the second day.

Results: Changes in nectar guide colour in A. szechenyi were intrinsic and age-dependent, although pollination somewhat accelerated the change. By the time that the nectar guides disappeared completely, floral rewards were reduced almost to zero. Artificial removal of nectar guides decreased both fruit set and pollen export. Flowers without nectar guides do not appear to increase the overall attractiveness of the plants.

Conclusions: Nectar guides and their changes represent reliable signals to pollinators and enhance both male and female reproductive success.     

Optimal pollinator attraction strategies in Trollius ranunculoides Hemsl. (Ranunculaceae) at different altitudes: increased floral display or promotion of nectar output?

For alpine plant species, patterns of resource allocation to functional floral traits for pollinator attraction can be highly significant in adaptation to low pollinator abundance and consequent pollen limitation. Increased pollination can be achieved either through a larger floral display or production of more pollen rewards. In this study, variation in resource allocation to different components for pollinator attraction was studied along an altitudinal gradient in Trollius ranunculoides, an obligate self‐incompatible out‐crosser of the Qinghai–Tibet Plateau. We compared resource allocation to conspicuous yellow sepals (which mainly provide visual attraction) and degenerate petals (which provide the major nectar reward) between populations at four altitudes. Furthermore, we investigated the contribution of sepals and petals to pollinator attraction and female reproductive success in an experiment with sepal or petal removal at sites at different altitudes. At the level of single flowers, resource allocation increased to sepals but decreased to petals with increasing altitude. Consistent with these results, sepals contributed much more to visitation rate and seed set than petals, as confirmed in the sepal or petal removal experiment. Sepals and petals contributed to female reproductive success by ensuring visitation rate rather than visitation duration. To alleviate increasing pollen limitation with increasing altitude, resource allocation patterns of T. ranunculoides altered to favour development of sepals rather than petals. This strategy may improve pollination and reproductive success through visual attraction (sepal) rather than nectar reward (petal) over a gradient of decreasing pollinator abundance.     

The dynamics of pollen removal and deposition,and its effects on sexual phases in a protandrous plant: Glechoma longituba (Lamiaceae)

The duration of sexual phases in dichogamous plants are affected by many factors. Using both experimental and observational studies, we investigated natural patterns of pollen removal and deposition, visiting frequency of pollinators, patterns of nectar secretion, and effects of pollen removal and stigmatic pollen deposition on the duration of sexual phases in a protandrous plant, Glechoma longituba. We found that visiting frequency of pollinators correlated with the nectar secretion pattern. The nectar volume during the male phase was higher than during the female phase. In the morning, the main pollinator, the bee Anthophora plumipes, mainly foraged for nectar and showed no preference for flowers in male or female phase, despite male phase flowers producing higher amounts of nectar. However, in the afternoon, they changed their behavior and foraged mainly for pollen, and then showed a preference for flowers in male phase. Furthermore, the rates of pollen removal and stigmatic pollen deposition can affect the starting time and the duration of the female phase. When pollen removal and pollination rates are low due to scarcity of pollinator services, the sexual phase can be prolonged, leading to an overlap, and thereby enhance the chance for sexual reproduction through pollinator‐facilitated self‐pollination. We consider the variation of sexual phases in Glechoma longituba an adaptive mechanism prepared for both cross‐pollination enhancement and reproductive assurance depending on the available pollination services.     

Novel nectar robbing negatively affects reproduction in Digitalis purpurea

With many plant–pollinator interactions undergoing change as species’ distributions shift, we require a better understanding of how the addition of new interacting partners can affect plant reproduction. One such group of floral visitors, nectar robbers, can deplete plants of nectar rewards without contributing to pollination. The addition of nectar robbing to the floral visitor assemblage could therefore have costs to the plant´s reproductive output. We focus on a recent plant colonist, Digitalis purpurea, a plant that in its native range is rarely robbed, but experiences intense nectar robbing in areas it has been introduced to. Here, we test the costs to reproduction following experimental nectar robbing. To identify any changes in the behavior of the principal pollinators in response to nectar robbing, we measured visitation rates, visit duration, proportion of flowers visited, and rate of rejection of inflorescences. To find the effects of robbing on fitness, we used proxies for female and male components of reproductive output, by measuring the seeds produced per fruit and the pollen export, respectively. Nectar robbing significantly reduced the rate of visitation and lengths of visits by bumblebees. Additionally, bumblebees visited a lower proportion of flowers on an inflorescence that had robbed flowers. We found that flowers in the robbed treatment produced significantly fewer seeds per fruit on average but did not export fewer pollen grains. Our finding that robbing leads to reduced seed production could be due to fewer and shorter visits to flowers leading to less effective pollination. We discuss the potential consequences of new pollinator environments, such as exposure to nectar robbing, for plant reproduction.     

Functional implications of the specialized staminal appendages in alpine ginger (Roscoea spp.: Zingiberaceae)

Floral organs are widely believed to enhance the pollination and reproductive success of angiosperms. However, the functional implication of some floral structures is still unknown. In this study, we explored the functional role of staminal appendages on male and female reproductive success of Himalayan Roscoea spp. and tested if their function differed between species with biotic pollination and autonomous selfing. Phenotypic manipulation is a powerful approach to test the functional effect of a particular trait on plant fitness. We compared various proxies of pollination success between intact flowers and flowers with manually excised staminal appendages. We found that the rate of visitation did not differ between intact and manipulated flowers. Our results revealed that in outcrossing Roscoea spp., the staminal appendages act as triggering devices to facilitate pollen release and deposition and also to manipulate the foraging position of pollinators to ensure both male and female reproductive success. In contrast, in autonomously selfing Roscoea spp., the removal of staminal appendages did not affect any aspect of pollination processes. Our results suggest that the staminal appendages are an integral component of outcrossing in Roscoea spp. and are maintained by selection pressure through both male and female reproductive success. This study provides important insights on how variation in breeding systems can provoke changes in the structure and function of floral organs among congeners.     

Effects of floral herbivory, limited pollination, and intrinsic plant characteristics on phenotypic gender in Sanicula arctopoides

Sexual expression in hermaphroditic plants is often a function of environmental factors affecting individuals before or during flowering. I tested for the effects of floral herbivory and lack of pollination in early umbels on the relative proportions of hermaphroditic and staminate (male) flowers produced on later umbels by Sanicula arctopoides, a monocarpic, andromonoecious perennial. Neither floral herbivory or lack of early pollination had a significant effect on the ratio of the two floral morphs, but the probability of producing staminate flowers on late umbels was strongly and positively related to plant size measured just prior to floral initiation and prior to herbivory. Plant size was also negatively correlated with flowering date. I suggest that producing staminate flowers on late umbels should benefit large early-blooming plants more than small late-blooming plants because more mating opportunities occur during the period when these flowers release pollen. Although herbivory did not cause labile changes of sex, whole plant phenotypic gender was still strongly affected by various forms of treatment. Sex-biased herbivory or lack of pollination rendered plants more or less phenotypically male, depending on which tissues were affected. Deer and pollen-feeding mites preferentially remove male tissues while hymenopteran seed predators preferentially remove female tissues. I conclude that combinations of herbivores could have counteracting or compounding effects on plant gender, and these effects may change the rankings of male and female reproductive success within populations. Received: 20 February 1996/Accepted: 30 July 1996