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.     

Massive floral display affects insect visits but not pollinator‐mediated pollen transfer in Rhododendron ferrugineum

Fragmentation of natural vegetation creates one of the largest threats to plant–pollinator interactions. Although fragmentation impacts on plant populations have been explored in many, mainly herbaceous, species, the response of wild mass‐flowering species is poorly known. Here, we studied 28 heathland patches dominated by the mass‐flowering shrub Rhododendron ferrugineum, each presenting different R. ferrugineum floral display sizes (total inflorescence number per patch) and patch isolation (median distance to the three nearest patches). We assessed the impacts of these two factors on (i) heathland patch visitor assemblage (considering R. ferrugineum versus surrounding community) and (ii) R. ferrugineum flower visitation rate and pollen transfer limitation (comparing seed set from emasculated to pollen‐supplemented flowers). We found that diversity and abundance of bees visiting R. ferrugineum in heathland patches significantly decreased with decreasing R. ferrugineum floral display, while overall visitor density per patch and flower visitation rate increased. Moreover, a decrease in massive floral display and increase in patch isolation resulted in reduced visitor density in the surrounding community. Even in patches with few individuals, we found disproportionate visitor abundance in R. ferrugineum compared to the surrounding community. Finally, pollen transfer limitation in R. ferrugineum was neither affected by visitation rate nor by patch attributes. By disproportionally attracting pollinators from co‐flowering species, and probably promoting geitonogamous pollen transfer, the mass‐flowering trait appears adequate to compensate, in terms of conspecific pollen transfer, for the decrease in visitor diversity and abundance and in mate availability, which usually result from population fragmentation.     

High invasive pollen transfer, yet low deposition on native stigmas in a Carpobrotus-invaded community

Background and Aims

Invasive plants are potential agents of disruption in plant–pollinator interactions. They may affect pollinator visitation rates to native plants and modify the plant–pollinator interaction network. However, there is little information about the extent to which invasive pollen is incorporated into the pollination network and about the rates of invasive pollen deposition on the stigmas of native plants.

Methods

The degree of pollinator sharing between the invasive plant Carpobrotus affine acinaciformis and the main co-flowering native plants was tested in a Mediterranean coastal shrubland. Pollen loads were identified from the bodies of the ten most common pollinator species and stigmatic pollen deposition in the five most common native plant species.

Key Results

It was found that pollinators visited Carpobrotus extensively. Seventy-three per cent of pollinator specimens collected on native plants carried Carpobrotus pollen. On average 23 % of the pollen on the bodies of pollinators visiting native plants was Carpobrotus. However, most of the pollen found on the body of pollinators belonged to the species on which they were collected. Similarly, most pollen on native plant stigmas was conspecific. Invasive pollen was present on native plant stigmas, but in low quantity.

Conclusions

Carpobrotus is highly integrated in the pollen transport network. However, the plant-pollination network in the invaded community seems to be sufficiently robust to withstand the impacts of the presence of alien pollen on native plant pollination, as shown by the low levels of heterospecific pollen deposition on native stigmas. Several mechanisms are discussed for the low invasive pollen deposition on native stigmas.Key words: Alien plant, Carpobrotus aff. acinaciformis, competition for pollinators, invasion, Mediterranean shrubland, plant-pollinator network, pollen loads, pollinator visits, stigma     

塔落岩黄芪主要传粉蜂的传粉效率研究

塔落岩黄芪 (Hedysarum laeve)是毛乌素沙地重要豆科植物, 在我国西部生态系统恢复与重建中起着重要作用, 但其自交率非常低, 必须依靠传粉昆虫授粉才能结实。野生传粉蜂是其主要传粉者。为了明确塔落岩黄芪的主要传粉蜂组成及优势传粉蜂的传粉效率, 2004–2006年, 我们设置了4个2 m×2 m样方观测访花昆虫的组成并对主要传粉蜂的访花频率、花粉移出率和柱头花粉沉降数目及单花停留时间进行了观测。结果表明白脸条蜂(Anthophora albifronella)、海切叶蜂(Megachile maritima)和散熊蜂(Bombus sporsdicus)是塔落岩黄芪的主要传粉蜂, 其中白脸条蜂在数量和访花频率上占有明显优势, 但其花粉移出率比海切叶蜂和散熊蜂的低, 3种蜂的柱头花粉沉降数目没有显著区别。通过对这3种蜂的花粉移出率、柱头花粉沉降数目和访花频率综合分析后, 我们认为白脸条蜂是塔落岩黄芪最有效的传粉蜂。     

Effects of pollen reward removal on fecundity in a self-incompatible hermaphrodite plant

The pollen of hermaphrodite plants is often utilised by flower-visiting animals. While pollen production has obvious benefits for plant male fitness, its consequences for plant female fitness, especially in self-incompatible hermaphrodite species, are less certain. Pollen production could either enhance seed production though increased pollinator attraction, or reduce it if ovules are discounted by deposition of self pollen, as can occur in species with late-acting self-incompatibility. To test the effects of pollen reward provision on female fitness, we artificially emasculated flowers in two populations of the succulent Aloe maculata (Asphodelaceae), which has a late-acting self-incompatibility system, over the course of its flowering period. Flowers of this species are visited by sunbirds (for nectar) and native bees (for pollen and nectar). We measured floral visitation rates, floral rejection rates, pollen deposition on stigmas and fruit and seed set in both emasculated and non-emasculated plants. We found that flowers of emasculated plants suffered reduced visitation and increased rejection (arrival without visitation) by bees, but not by sunbirds; had fewer pollen grains deposited on stigmas and showed an overall decrease in fruit set and seed set. Rates of seed abortion were, however, greatly reduced in emasculated flowers. This study shows that pollen rewards can be important for seed set, even in self-incompatible plants, which have been assumed to rely on nectar rewards for pollinator attraction. Seed abortion was, however, increased by pollen production, a result that highlights the complexity of selection on pollen production in hermaphrodite flowers.     

The reproductive assurance benefit of selfing: importance of flower size and population size

Autonomous selfing can provide reproductive assurance (RA) for flowering plants that are unattractive to pollinators or in environments that are pollen limited. Pollen limitation may result from the breakdown of once-continuous habitat into smaller, more isolated patches (habitat fragmentation) if fragmentation negatively impacts pollinator populations. Here we quantify the levels of pollen limitation and RA among large and small populations of Collinsia parviflora, a wildflower with inter-population variation in flower size. We found that none of the populations were pollen limited, as pollen-supplemented and intact flowers did not differ in seed production. There was a significant effect of flower size on RA; intact flowers (can self) produced significantly more seeds than emasculated flowers (require pollen delivery) in small-flowered plants but not large-flowered plants. Population size nested within flower size did not significantly affect RA, but there was a large difference between our two replicate populations for large-flowered, small populations and small-flowered, large populations that appears related to a more variable pollination environment under these conditions. In fact, levels of RA were strongly negatively correlated with rates of pollinator visitation, whereby infrequent visitation by pollinators yielded high levels of RA via autonomous selfing, but there was no benefit of autonomous selfing when visitation rates were high. These results suggest that autonomous selfing may be adaptive in fragmented habitats or other ecological circumstances that affect pollinator visitation rates.     

Change of floral orientation affects pollinator diversity and their relative importance in an alpine plant with generalized pollination system,Geranium refractum (Geraniaceae)

Floral orientation may affect pollinator attraction and pollination effectiveness, and its influences may differ among pollinator species. We, therefore, hypothesized that, for plant species with a generalized pollination system, changes in floral orientation would affect the composition of pollinators and their relative contribution to pollination. Geranium refractum, an alpine plant with downward floral orientation was used in this study. We created upward-facing flowers by altering the flower angle. We compared the pollinator diversity, pollination effectiveness, and pollinator importance, as well as female reproductive success between flowers with downward- and upward-facing orientation. Results indicated that the upward-facing flowers were visited by a wider spectrum of pollinators (classified into functional groups), with higher pollinator diversity than natural flowers. Moreover, due to influences on visitation number and pollen removal, the pollinator importance exhibited by the main pollinator groups differed between flower types. Compared with natural flowers, the pollination contribution of principal pollinators (i.e., bumblebees) decreased in upward-facing flowers and other infrequent pollinators, such as solitary bees and muscoid flies, removed more pollen. Consequently, stigmatic pollen loads were lower in upward- than in downward-facing flowers. These findings reveal that floral orientation may affect the level of generalization of a pollination system and the relative importance of diverse pollinators. In this species, the natural downward-facing floral orientation may increase pollen transfer by effective pollinators and reduce interference by inferior pollinators.     

Pollination efficiency of cockroaches and other floral visitors of Clusia blattophila

• Cockroaches have rarely been documented as pollinators. In this paper we examine whether this is because they might be inefficient at pollination compared to other pollinators. Clusia blattophila, a dioecious shrub growing on isolated rocky outcrops in French Guiana, is pollinated by Amazonina platystylata cockroaches and provides a valuable system for the study of cockroach pollination efficiency.
• We examined the species composition of the visitor guild and visitation rates by means of camcorder recordings and visitor sampling. Then, we investigated the capacity for pollen transfer of principal visitors and found correlations between visitation rates and pollen loads on stigmas. In an exclusion experiment we determined the contributions of individual species to pollination success.
• Amazonina platystylata, crickets and two species of Diptera transferred pollen, but the number of transferred pollen grains was only related to visitation rates in the case of cockroaches. Crickets visited and rarely carried pollen. Dipterans were as frequent as cockroaches, carried similar pollen loads, but transferred much less pollen. An estimated 41% and 17% of ovules were pollinated by cockroaches and dipterans, respectively. The remaining ovules were not pollinated. There was no spatial variation in pollinator guild composition, but cockroaches visited flowers less frequently at the smaller study site.
• We demonstrate that cockroaches pollinate a large proportion of ovules. Their pollination service is not confined to one study site and, unlike that provided by dipterans, is not limited to certain years. We suggest that cockroach pollination has been overlooked and that cockroach‐pollinated plants, which share certain floral features, possess adaptations to pollination by cockroaches.

     

Extremely high proportions of male flowers and geographic variation in floral ratios within male figs of Ficus tikoua despite pollinators displaying active pollen collection

Most plants are pollinated passively, but active pollination has evolved among insects that depend on ovule fertilization for larval development. Anther‐to‐ovule ratios (A/O ratios, a coarse indicator of pollen‐to‐ovule ratios) are strong indicators of pollination mode in fig trees and are consistent within most species. However, unusually high values and high variation of A/O ratios (0.096–10.0) were detected among male plants from 41 natural populations of Ficus tikoua in China. Higher proportions of male (staminate) flowers were associated with a change in their distribution within the figs, from circum‐ostiolar to scattered. Plants bearing figs with ostiolar or scattered male flowers were geographically separated, with scattered male flowers found mainly on the Yungui Plateau in the southwest of our sample area. The A/O ratios of most F. tikoua figs were indicative of passive pollination, but its Ceratosolen fig wasp pollinator actively loads pollen into its pollen pockets. Additional pollen was also carried on their body surface and pollinators emerging from scattered‐flower figs had more surface pollen. Large amounts of pollen grains on the insects' body surface are usually indicative of a passive pollinator. This is the first recorded case of an actively pollinated Ficus species producing large amounts of pollen. Overall high A/O ratios, particularly in some populations, in combination with actively pollinating pollinators, may reflect a response by the plant to insufficient quantities of pollen transported in the wasps’ pollen pockets, together with geographic variation in this pollen limitation. This suggests an unstable scenario that could lead to eventual loss of wasp active pollination behavior.     

FLORAL DISPLAY IN PHYLA INCISA: CONSEQUENCES FOR MALE AND FEMALE REPRODUCTIVE SUCCESS

The inflorescences of Phyla incisa consist of flowers in two phases: younger, nectar-containing flowers that have yellow corolla throats and older, nectar-lacking flowers that have dark purple corolla throats. Observations of pollinator visitation patterns to both natural and manipulated inflorescences were made to determine the role of each flower phase in pollinator attraction. The effect of older-phase flowers on male and female reproductive success was determined by comparing stigmatic pollen loads and estimates of pollen removal from inflorescences having different numbers of these flowers. The pollinators of Phyla selected larger inflorescences more often than expected based upon the size distribution of inflorescences available to them. Both younger- and older-phase flowers contributed to the attraction of pollinators, but the latter were less effective in this function. The presence of older-phase flowers significantly increased the visitation rate to inflorescences and the amount of pollen removed but had little effect on pollen deposition on stigmas. The lack of correspondence between pollen deposition and pollinator-visitation rate was not due to stigma saturation, since stigma loads varied greatly. The data indicate that the deposition of pollen on stigmas in this species is a relatively stochastic process, whereas pollen removal from inflorescences occurs at a much more regular rate. Old-phase flower retention appears to contribute to reproductive success through increased pollen donation when pollinator activity is high and may also increase the probability of seed set when pollinators are rare.     

Seasonal and annual variations in the pollination efficiency of a pollinator community of Dictamnus albus L.

The interplay between insect and plant traits outlines the patterns of pollen transfer and the subsequent plant reproductive fitness. We studied the factors that affect the pollination efficiency of a pollinator community of Dictamnus albus L. by evaluating insect behaviour and morphological characteristics in relation to flowering phenology. In order to extrapolate the pollinator importance of single taxa and of the whole pollinator guild, we calculated an index distinguishing between potential (PPI) and realized (RPI) pollinator importance. Although the pollinator species spectrum appeared rather constant, we found high intra‐ and inter‐annual variability of pollinator frequency and importance within the insect community. Flower visitation rate strictly depended on insect abundance and on the overlap between their flying period and flower blooming. All the pollinators visited flowers from the bottom to the top of the racemes, excluding intra‐plant geitonogamous pollination, and most of them showed high pollen fidelity. Only medium large‐sized bees could contact the upward bending stiles while feeding on nectar, highlighting a specialisation of the plant towards bigger pollinators. Moreover, we found evidence of functional specialisation, since all pollinators were restricted to a single taxonomic group (order: Hymenoptera; superfamily: Apoidea). Both the PPI and RPI indices indicate Habropoda tarsata as the most important pollinator of D. albus. Following hand cross‐pollination experiments we revealed the presence of pollination limitation in 1 of the 3 years of field study. We discuss this result in relation to flowering abundance and to possible mismatches of phenological periods between plants and insects.     

Effectiveness of buzz pollination in Pedicularis chamissonis: significance of multiple visits by bumblebees

Pollination success of plants is highly susceptible to the frequency of visits and foraging behavior of pollinators. Pollination of the nectarless flowers of Pedicularis species depends on bumblebee workers collecting pollen by vibrating the anthers (buzz pollination). However, little is known about the efficiency of the pollination system. Foraging behavior, pollen removal from anthers and pollen deposition on stigmas of P. chamissonis were studied to assess the effectiveness of buzz pollination in an alpine snowbed population of northern Japan. Although bumblebees tended to visit most of the flowers open at a given time within inflorescences during a single visit, pollen removal rate at the first visit was about 20%, and buzzing period decreased with increasing number of previous visits, resulting in a decreasing proportion of pollen removed per visit as the number of visits increased. These trends enable plants to provide pollen for more pollinators. The number of pollen grains deposited on stigmas was not saturated during the first visit and increased with additional visits. Irrespective of weak self-compatibility, evidence of interference between self and outcross pollen was lacking for seed production. Therefore, buzz pollination in P. chamissonis acts as a mechanism that improves the chance of cross-pollination upon multiple visits if pollinator visitation is frequent.     

The relative importance of solitary bees and syrphid flies as pollinators of two outcrossing plant species in the New Zealand alpine

Pollinators vary in their relative contribution to the conspecific pollen deposited onto receptive stigmas, because of variation in both visitation rate and effectiveness of pollen transfer. Syrphid flies and short‐tongued solitary bees are common flower visitors in alpine New Zealand, yet their relative importance as pollinators is unknown. We measured pollinator performance of the New Zealand alpine endemics Hylaeus matamoko (Hymenoptera: Colletidae) and Allograpta spp. (Diptera: Syrphidae) on two New Zealand alpine herbs, Ourisia glandulosa (Plantaginaceae) and Wahlenbergia albomarginata (Campanulaceae). Ourisia glandulosa received visits by solitary bees and syrphid flies at equal frequencies, whereas W. albomarginata was mostly visited by H. matamoko. Based on single‐visit pollen deposition to virgin stigmas, H. matamoko was a much more effective pollinator than Allograpta spp., delivering 10 times as much pollen per visit to O. glandulosa stigmas and 3 times as much to W. albomarginata stigmas. By multiplying visitation frequency by single‐visit pollen deposition, we estimated that H. matamoko performed 90% and 95% of the pollination of O. glandulosa and W. albomarginata, respectively. Although H. matamoko bees are short‐tongued and small in size, they are critically important to plant reproductive success in the New Zealand alpine. These bees contributed most of the pollination, even to a species that received just as many visits by flies, underscoring the need to consider per‐visit effectiveness as well as visitation rate in assessing the importance of different pollinators.     

One for all and all for one: retention of colour‐unchanged old flowers increases pollinator attraction in a hermaphroditic plant

• Long‐lived flowers increase pollen transfer rates, but these entail high water and carbon maintenance costs. The retention of pollinated and reward‐free old flowers enhances pollinator visitation to young receptive flowers by increasing floral display size. This mechanism is associated with acropetal inflorescences or changes in flower colour and openness, but the retention of unchanging solitary flowers remains overlooked.
• We examined pollination‐dependent variation in floral longevity and determined stigmatic receptivity, pollen viability and pollen removal rates among flower ages in Kielmeyera regalis, a Neotropical savanna shrub. We also evaluated the effects of floral display size on pollinator visitation rates. Lastly, we determined whether old flowers are unvisited and exclusively increase pollinator attraction to young flowers through flower removal experiments.
• Regardless of pollination treatment, flowers lasted fully open with no detectable physical changes for 3 days. Over time, stigmas remained receptive but >95% of pollen was removed. Pollinator visitation significantly increased with floral display size and intermediate percentages (15–30%) of newly opened flowers. Accordingly, the retention of reward‐free and unvisited old flowers increased young flower–pollinator interaction.
• Our results reveal the importance of a prolonged floral longevity in increasing pollinator attraction toward newly opened receptive flowers without changes in flower colour and form. We conclude that the retention of pollinated, reward‐free and unvisited colour‐unchanged old flowers in K. regalis is a strategy that counteracts the water use costs associated with the maintenance of large flowers with increased mate opportunities in a pollen‐limited scenario.

     

Effects of flowering plant density on pollinator visitation, pollen receipt, and seed production in Delphinium barbeyi (Ranunculaceae)

Variation in flowering plant density can have conflicting effects on pollination and seed production. Dense flower patches may attract more pollinators, but flowers in those patches may also compete for pollinator visits and abiotic resources. We examined how natural and experimental conspecific flowering plant density affected pollen receipt and seed production in a protandrous, bumble bee-pollinated wildflower, Delphinium barbeyi (Ranunculaceae). We also compared floral sex ratios, pollinator visitation rates, and pollen limitation of seed set from early to late in the season to determine whether these factors mirrored seasonal changes in pollen receipt and seed production. Pollen receipt increased with natural flowering plant density, while seed production increased across lower densities and decreased across higher flower densities. Experimental manipulation of flowering plant density did not affect pollinator visitation rate, pollen receipt, or seed production. Although pollinator visitation rate increased 10-fold from early to late in the season, pollen receipt and seed set decreased over the season. Seed set was never pollen-limited. Thus, despite widespread effects of flowering plant density on plant reproduction in other species, the effects of conspecific flowering plant density on D. barbeyi pollination and seed production are minor.     

Pollination ecology of a plant in its native and introduced areas

Entomophilous and obligate out-crossing non-native plants need to become well integrated in the resident plant–pollinator network to set seeds and become established. However, it is largely unknown how pollination patterns differ between native ranges and those where plants have been introduced.We compared the identity, abundance and visitation rates of pollinators, insect pollen loads, pollen deposition on stigmas, and fruit and seed sets of Hedysarum coronarium, an entomophilous short lived N-fixing perennial, in populations from native and introduced ranges in Spain (South of mainland Spain and Menorca Island, respectively).In both areas, Hedysarum was visited by a similar number of species, mainly hymenopterans; seven species were common between native and introduced areas. However, pollinator richness, abundance, and visits per flower were greater in the native than in the introduced range, as were fruit and seed sets. Hedysarum pollen loads on stigmas and on Apis mellifera, the most common pollinator, did not differ between areas. Lower abundance of pollinators might be causing lower visitation rates, and to some extent reducing Hedysarum fruit and seed sets in the introduced area.Our biogeographical approach shows that integration of a non-native plant in a resident pollinator network does not prevent pollen limitation in the introduced area. Therefore, despite being necessary, pollination mutualistic relationships might not be the key for non-native plant establishment success in the introduced area.     

铃铛子和赛莨菪（茄科）的传粉机制比较研究

通过野外交配系统试验和传粉昆虫观察,以铃铛子（Anisodus luridus）和赛莨菪（A．camiolicoides）为研究对象,探讨了山莨菪属内自交亲和系统的进化与传粉昆虫的转变。结果表明,铃铛子和赛莨菪均属于自交完全亲和的类群,但两个种的自动自交能力均不强,而且都存在传粉限制。北方黄胡蜂（Vespula rtfarufa）和石长黄胡蜂（Dolichovespula saxonica）分别是铃铛子和赛莨菪的主要传粉昆虫,自然状态下两种昆虫的传粉效率均比较高,但两种昆虫访问两种植物时传递花粉的方式不同,其中北方黄胡蜂主要促进了铃铛子的花间传粉,而石长黄胡蜂访花引起赛莨菪同一朵花内的传粉。通过与该属的另一种植物山莨菪（A．tanguticus）的传粉机制比较,发现在山莨菪属的物种分化过程中,由自交不亲和转变为自交完全亲和,传粉昆虫也发生了转变,证明了自交亲和系统存在于起源较晚的类群中。高山环境中频繁且不可预测的降雨可能降低了传粉昆虫的活动能力,进而导致铃铛子和赛莨菪均存在传粉限制。     

THE EFFECT OF DELPHINIUM NELSONII POLLEN ON SEED SET IN IPOMOPSIS AGGREGATA,A COMPETITOR FOR HUMMINGBIRD POLLINATION

Sympatric plant species can compete for pollination services in several ways. For example, pollinators may move between species and deposit heterospecific pollen on stigmas, which in turn may reduce the efficacy of conspecific pollen. We explored this possibility by determining the effect of Delphinium nelsonii pollen on seed set in Ipomopsis aggregata. These montane herbs are pollinated by hummingbirds, experience heterospecific pollen deposition in nature, and suffer reduced seed set in each other's presence. We hand-pollinated flowers of I. aggregata with either pure conspecific pollen or a mixture of pollen of the two species. Resulting pollen loads on stigmas ranged from 0–865 D. nelsonii grains and from 10–336 I. aggregata grains; mean seed set per flower was 11.3. There was no detectable effect of D. nelsonii pollen load on I. aggregata seed set. It is possible that seed set reductions seen in previous studies of competition for pollination between these species were caused by pollen wastage, pollen layering on the pollinator, or the temporal sequence of pollen arrival at the stigma.     

The plot thickens: does low density affect visitation and reproductive success in a perennial herb, and are these effects altered in the presence of a co-flowering species?

Plants may experience reduced reproductive success at low densities, due to lower numbers of pollinator visits or reduced visit quality. Co-occurring plant species that share pollinators have the potential to facilitate pollination by either increasing numbers of pollinator visits or increasing the quality of visits, but also have the potential to reduce plant reproductive success through competition for pollination. I used a field experiment with a common distylous perennial (Piriqueta caroliniana) in the presence and absence of a co-flowering species (Coreopsis leavenworthii) in plots with one of four different distances between conspecific plants. I found strong negative effects of increasing interplant distance (related to conspecific density) on several components of P. caroliniana reproductive success: pollinator visits to plants per plot visit, visits received by individual plants, conspecific pollen grains on stigmas, outcross pollen grains on stigmas, and probability of fruit production. Although P. caroliniana and C. leavenworthii share pollinators, the co-flowering species did not affect visitation, pollen receipt or reproductive effort in P. caroliniana. Pollinators moved very infrequently between species in this experiment, so floral constancy might explain the lack of effect of the co-flowering species on P. caroliniana reproductive success at low densities. In co-occurring self-incompatible plants with floral rewards, reproductive success at low density may depend more on conspecific densities than on the presence of other species.     

FLORAL BIOLOGY,HUMMINGBIRD POLLINATION AND FRUIT PRODUCTION OF TRUMPET CREEPER (CAMPSIS RADICANS,BIGNONIACEAE)

Trumpet creeper is self-incompatible and bears long, tubular, orange flowers from June to September. Flowering peaks rapidly, then declines and continues at low levels for several weeks. The initial burst of flowering may attract pollinators that return even during subsequent reduced flowering. Most flowers open before noon and nectar production totals 110 μl of 26% sucrose equivalents per flower, an exceptionally high production for a temperate zone plant. Production ceases within 20–30 hr of flower opening, but corollas persist for several days and may serve to attract pollinators. Effective pollination reduces the period of stigma receptivity and speeds closing of stigma lobes. Only 1–9% of flowers produced mature fruits at four sites in Illinois and Missouri. Roughly 400 pollen grains had to be deposited on a receptive stigma to cause fruit development beyond an initial period of high abortion. At two sites, 17% and 89% of stigmas received over 400 pollen grains. Assuming 50% of deposited grains were from the same plant, fruit production at one site was clearly pollinator limited, that at the second site may have been. Ruby-throated Hummingbirds (Archilochus colubris) deposited ten times as much pollen per stigma per visit as honeybees (Apis mellifera) and bumblebees (Bombus spp.). Fruit set was highest where rubythroat visitation was most frequent. Trumpet creeper appears primarily adapted for hummingbird pollination, but can also be adequately pollinated by honeybees and bumblebees. This is one of the first attempts to relate pollen-depositing capabilities of pollinators of any plant to pollen requirements for fruit production. Several characteristics suggest that trumpet creeper may be adapted to pollination at low densities (often called traplining) in its presumed original, woodland, habitat.