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

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

Selection of trait combinations through bee and fly visitation to flowers of Polemonium foliosissimum

Pollinators are known to exert natural selection on floral traits, but the extent to which combinations of floral traits are subject to correlational selection (nonadditive effects of two traits on fitness) is not well understood. Over two years, we used phenotypic manipulations of plant traits to test for effects of flower colour, flower shape and their interaction on rates of pollinator visitation to Polemonium foliosissimum. We also tested for correlational selection based on weighting visitation by the amount of conspecific pollen delivered per visit by each category of insect visitor. Although bumblebees were the presumed pollinators, solitary bees and flies contributed substantially (42%) to pollination. In manipulations of one trait at a time, insects visited flowers presenting the natural colour and shape over flowers manipulated to present artificial mutants with either paler colour or a more open or more tubular flower. When both colour and shape were manipulated in combination, selection on both traits arose, with bumblebees responding mainly to colour and flies responding mainly to shape. Despite selection on both floral traits, in a year with many bumblebees, we saw no evidence for correlational selection of these traits. In a year when flies predominated, fly visitation showed a pattern of correlational selection, but not favouring the natural phenotype, and correlational selection was still not detected for expected pollen receipt. These results show that flower colour and shape are subject to pollinator‐mediated selection and that correlational selection can be generated based on pollinator visitation alone, but provide no evidence for correlational selection specifically for the current phenotype.     

Association of flower color with pollen reward may explain increased bumblebee visitation to the Scotch broom yellow morph

Given that pollinators usually visit flowers for hidden rewards, they need to rely on floral traits that indicate reward status (“honest signals”). However, the relationship between pollination, honest signals, and floral rewards is little documented in natural conditions. The Scotch broom (Cytisus scoparius) is an invasive shrub with polymorphism in the color of its flowers that can be yellow, orange, or red. In three areas dominated by the Scotch broom, we described the abundance of the floral morphs and estimated bumblebee (Bombus terrestris) visitation rate. We examined whether bumblebee visitation to the floral morphs was related to pollen reward. We collected flowers and classified their stamens according to their function: reward or pollen export. Then, we measured anther size and estimated pollen quantity. The yellow morph was more abundant and more visited by bumblebees than the orange and red morphs. The yellow flowers did indeed offer more pollen than the other morphs and this occurred only for rewarding anthers, suggesting that bumblebees could use yellow color as an honest signal to visit the most rewarding flowers. We discuss whether innate and/or learned preferences of bumblebees can explain why the yellow morph is more visited, pollinated, and abundant, while the other morphs are maintained at a lower frequency. This is one of the few field works that shows that variation in intra-specific floral traits is associated with variation in floral reward and pollinator visitation rate, helping to understand the foraging preferences of pollinators and the coexistence of floral morphs in nature.

Clinical trials registration: Not applicable.

     

Causes and consequences of floral damage in Aconitum lycoctonum at high and low elevations in Switzerland

The outcomes of interactions among plants and the insects that use their flowers are likely to vary among the physical environments and the communities in which they grow. In this study we quantified floral damage of Aconitum lycoctonum in high (>2000 m) and low (c. 500 m) elevation populations in Switzerland. At high elevation, floral damage was frequent and was caused by nectar-robbing short-tongued bumblebees. Nectar robbers make a hole in the flower when they collect nectar. A nectar robber exclusion experiment showed that nectar robbery by short-tongued bumblebees had no effect on the female reproductive success of plants; robbing bees rarely damaged the nectaries, and damage to the petals probably does not decrease flower longevity. In addition, nectar robbers tended to collect pollen during about 10% of their visits. Thus, these bees may act as low-efficiency pollinators and may, at times, be mutualistic associates. At low elevation, the holes in the flowers were caused by beetles (Meligethes viridescens) and not by short-tongued bumblebees. The beetles eat pollen and might also consume nectar. Since the beetles gain access to pollen and nectar by entering the flower through its opening, and later chew holes while foraging on floral tissue, the beetles are pollen eaters rather than nectar robbers. Overall, our results show that not all floral damage is caused by nectar robbers and that there can be strong altitudinal variation in the causes and consequences of floral damage.     

Bombus terrestris in a mass‐flowering pollinator‐dependent crop: A mutualistic relationship?

Bumblebees (Bombus spp.) rely on an abundant and diverse selection of floral resources to meet their nutritional requirements. In farmed landscapes, mass‐flowering crops can provide an important forage resource for bumblebees, with increased visitation from bumblebees into mass‐flowering crops having an additional benefit to growers who require pollination services. This study explores the mutualistic relationship between Bombus terrestris L. (buff‐tailed bumblebee), a common species in European farmland, and the mass‐flowering crop courgette (Cucurbita pepo L.) to see how effective B. terrestris is at pollinating courgette and in return how courgette may affect B. terrestris colony dynamics. By combining empirical data on nectar and pollen availability with model simulations using the novel bumblebee model Bumble‐BEEHAVE, we were able to quantify and simulate for the first time, the importance of courgette as a mass‐flowering forage resource for bumblebees. Courgette provides vast quantities of nectar to ensure a high visitation rate, which combined with abundant pollen grains, enables B. terrestris to have a high pollination potential. While B. terrestris showed a strong fidelity to courgette flowers for nectar, courgette pollen was not found in any pollen loads from returning foragers. Nonetheless, model simulations showed that early season courgette (nectar) increased the number of hibernating queens, colonies, and adult workers in the modeled landscapes. Synthesis and applications. Courgette has the potential to improve bumblebee population dynamics; however, the lack of evidence of the bees collecting courgette pollen in this study suggests that bees can only benefit from this transient nectar source if alternative floral resources, particularly pollen, are also available to fulfill bees’ nutritional requirements in space and time. Therefore, providing additional forage resources could simultaneously improve pollination services and bumblebee populations.     

The floral biology ofThelymitra epipactoides (Orchidaceae), and the implications of pollination by deceit on the survival of this rare orchid

Thelymitra epipactoides has a highly variable visual display achieved through polychromatic flowers and variable inflorescence size, bearing between 7 and 31 flowers, which attract foraging polylectic bees. Only bees of the genusNomia were observed carrying pollinia and successfully pollinating the orchid. The genusNomia contains polylectic, pollen gathering species that store pollen in both the crop and scopa on the hind legs. The absence of a reward for the bees indicates the orchid is relying on deception to attract visitors. The relationship of deception to mimicry is discussed. Once on the flower, tactile, visual and possibly olfactory stimuli direct bees to the false anther formed by the voluminous column wings, where morphological adaptations of the flower ensure that the pollinarium is deposited on the gaster of the bee to effect pollination. — The lack of seed set observed on the Victorian coast appears to be due to the absence of pollinators from the heath and grassland communities in which the orchid grows. This may well be a consequence of the reduced number of plants flowering in the community (a result of the elimination of fire at these sites), thus not maintaining a floral community attractive to potential pollinators.     

The effects of landscape on bumblebees to ensure crop pollination in the highland agricultural ecosystems in China

Honey bees and wild bees provide critical pollination services to agricultural ecosystems; however, the relative contributions of different bee taxa are not well understood. The natural habitats surrounding farmland support food and nesting resources for wild bees and therefore play an important role in the maintenance of crop pollination. In this study, we selected Cucurbita pepo L. (squash) as a model crop to investigate the relative importance of honey bees and bumblebees in pollinating the crop. Thirteen fields, which were surrounded by a gradient of natural habitat, were investigated on the Yunnan‐Guizhou Plateau in China. We measured the visit densities of honey bees and bumblebees, the number of pollen grains deposited in a single visit by the two bee taxa, as well as the overall pollen grains deposited on stigmas during a flowering day, and then used Bayesian inference to decouple the pollen grains deposited by either the honey bees or the bumblebees. Compared with honey bees, bumblebees deposited a higher number of pollen grains on stigmas in a single visit, but had a lower visit density than honey bees. Meanwhile, the bumblebee visit density increased along the proportion of natural habitat, while the honey bee visit density was not affected by the surrounding natural habitat. Data simulations using Bayesian inference showed that on a flowering day, the number of pollen grains deposited by bumblebees increased with the proportion of natural habitat in the surrounding landscape, but the number of pollen grains deposited by honey bees did not. Moreover, the total numbers of pollen grains deposited by honey bees or bumblebees alone were all below 2000 (the critical level to satisfy the pollination requirement of this crop). Pollen calculations demonstrated that the number of pollen grains deposited by the two bee taxa was greater than 2000 in fields surrounded by more than 13% natural habitat (grasslands and forests). The results revealed that bumblebees ensured C. pepo pollination in combination with honey bees in the highland agricultural ecosystems.     

Bee visitation rates to cultivated sunflowers increase with the amount and accessibility of nectar sugars

Pollinators make foraging decisions based on numerous floral traits, including nectar and pollen rewards, and associated visual and olfactory cues. For insect‐pollinated crops, identifying and breeding for attractive floral traits may increase yields. In this study, we examined floral trait variation within cultivated sunflowers and its effects on bee foraging behaviours. Over 2 years, we planted different sunflower inbred lines, including male‐fertile and male‐sterile lines, and measured nectar volume, nectar sugar concentration and composition, and corolla length. During bloom, we recorded visits by both managed honey bees and wild bees. We then examined consistency in relative nectar production by comparing field results to those from a greenhouse experiment. Sunflower inbred lines varied significantly in all floral traits, including the amount and composition of nectar sugars, and in corolla length. Both wild bee and honey bee visits significantly increased with nectar sugar amount and decreased with corolla length, but appeared unaffected by nectar sugar composition. While wild bees made more visits to sunflowers providing pollen (male‐fertile), honey bees preferred plants without pollen (male‐sterile). Differences in nectar quantity among greenhouse‐grown sunflower lines were similar to those measured in the field, and bumble bees preferentially visited lines with more nectar in greenhouse observations. Our results show that sunflowers with greater quantities of nectar sugar and shorter corollas receive greater pollination services from both managed and wild bees. Selecting for these traits could thus increase sunflower crop yields and provide greater floral resources for bees.     

Colour learning when foraging for nectar and pollen: bees learn two colours at once

Bees are model organisms for the study of learning and memory, yet nearly all such research to date has used a single reward, nectar. Many bees collect both nectar (carbohydrates) and pollen (protein) on a single foraging bout, sometimes from different plant species. We tested whether individual bumblebees could learn colour associations with nectar and pollen rewards simultaneously in a foraging scenario where one floral type offered only nectar and the other only pollen. We found that bees readily learned multiple reward–colour associations, and when presented with novel floral targets generalized to colours similar to those trained for each reward type. These results expand the ecological significance of work on bee learning and raise new questions regarding the cognitive ecology of pollination.     

Nectar discovery speeds and multimodal displays: assessing nectar search times in bees with radiating and non-radiating guides

Floral displays are often composed of areas of contrasting stimuli which flower visitors use as guides, increasing both foraging efficiency and the likelihood of pollen transfer. Many aspects of how these displays benefit foraging efficiency are still unexplored, particularly those surrounding multimodal signals and the spatial arrangement of the display components. We compare the nectar discovery times of forager bumblebees (Bombus terrestris) when presented with artificial flowers with unimodal or compound displays of visual and/or olfactory stimuli, positioned in either radiating or non-radiating arrangements. We found that the addition of individual display components from either modality reduces nectar discovery time but there was no time benefit to bimodal displays over unimodal displays or any benefit to radiating stimuli arrangements over non-radiating arrangements. However, preference tests revealed a time advantage to radiating unimodal visual patterns over non-radiating unimodal visual patterns when both types were displayed simultaneously. These results suggest that the benefits of multimodal stimuli arrangements to pollinators are unrelated to benefits in nectar discovery time. Our results also suggest that spatial patterns of scent can be used as nectar guides and can reduce nectar discovery times without the aid of visual stimuli.     

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.     

Consumptive emasculation: the ecological and evolutionary consequences of pollen theft

Many of the diverse animals that consume floral rewards act as efficient pollinators; however, others 'steal' rewards without 'paying' for them by pollinating. In contrast to the extensive studies of the ecological and evolutionary consequences of nectar theft, pollen theft and its implications remain largely neglected, even though it affects plant reproduction more directly. Here we review existing studies of pollen theft and find that: (1) most pollen thieves pollinate other plant species, suggesting that theft generally arises from a mismatch between the flower and thief that precludes pollen deposition, (2) bees are the most commonly documented pollen thieves, and (3) the floral traits that typically facilitate pollen theft involve either spatial or temporal separation of sex function within flowers (herkogamy and dichogamy, respectively). Given that herkogamy and dichogamy occur commonly and that bees are globally the most important floral visitors, pollen theft is likely a greatly under-appreciated component of floral ecology and influence on floral evolution. We identify the mechanisms by which pollen theft can affect plant fitness, and review the evidence for theft-induced ecological effects, including pollen limitation. We then explore the consequences of pollen theft for the evolution of floral traits and sexual systems, and conclude by identifying key directions for future research.     

Selection for pollination success and the mechanical fit of Imfiatiens flowers around bumblebee bodies

Selection on flowers has often been viewed as being particularly strict, constant, and responsible for species differences. Impatiens pallida and I . capensis flowers fit snugly around bees, leading one to expect a close relationship between floral morphology and pollination success. My studies on the amount of pollen removed from androecia and deposited on stigmas in single visits by bumblebees did not confirm this supposition. Trimming off parts of the floral vestibule with scissors and gluing in pleats had very little effect on die amount of pollen that bees moved. In reciprocal transfer experiments, flowers from different populations sometimes differed in the amount of pollen moved, but when the two species were compared in sympatry, pollen removal and deposition differed hardly at all. A comparison of the relationship between pollen movement and floral morphology among 15 populations showed that, although there was great heterogeneity in die amount of pollen moved, the observed differences were independent of floral morphology. None of this supports a belief in strong selection that fine-tunes the mechanical fit between bee and flower; selection for visitation success based on pollinator behaviour may have a much stronger influence on floral characters.     

A generalised mimicry system involving angiosperm flower colour, pollen and bumblebees’ innate colour preferences

Flower colour is a major advertisement signal of zoophilous plants for pollinators. Bees, the main pollinators, exhibit innate colour preferences, which have often been attributed to only one single floral colour, though most flowers display a pattern of two or several colours. The existing studies of floral colour patterns are mostly qualitative studies. Using a model of bee colour vision we quantitatively investigate two questions: whether or not component colours of floral colour patterns may mimic pollen signals, and whether or not bumblebees exhibit innate preferences for distinct parameters of naturally existing floral colour patterns. We analysed the spectral reflectances of 162 plant species with multicoloured flowers and inflorescences, distiniguishing between inner and outer colours of floral colour patterns irrespective of the particular structures so coloured.We found that:– The inner colour of radially symmetrical flowers and inflorescences and of zygomorphic flowers appears less diverse to bees than the peripheral colour.– The inner colour of most radial flowers and inflorescences as well as the inner colour of a large number of non-related zygomorphic flowers appears to bees to be very similar to that of pollen.– Bumblebees (Bombus terrestris) exhibit innate preferences for two-coloured over single-coloured dummy flowers in a spontaneous choice test.– Bumblebees exhibit innate preferences for dummy flowers with a large over those with a small centre area.– Bumblebees exhibit innate preferences for dummy flowers with a centre colour similar to that of pollen over those with another centre colour.Our findings support the hypotheses that the inner component of floral colour patterns could be interpreted as a generalised and little recognised form of mimicry of the colour of visually displayed pollen, that bumblebees exhibit innate preferences regarding colour and size parameters of floral colour patterns, and that these correspond to visually displayed pollen. These findings together suggest a prominent role of floral colour patterns in advertisement to and guidance of naive flower visitors.     

Carry-over effects of bumblebee associative learning in changing plant communities leads to increased costs of foraging

Flower visitors learn to avoid food-deceptive plants and to prefer rewarding ones by associating floral cues to rewards. As co-occurring plant species have different phenologies, cue-reward associations vary over time. It is not known how these variations affect flower visitors’ foraging costs and learning. We trained bumblebees of two colonies to forage in a community of deceptive and rewarding artificial inflorescences whose flower colours were either similar or dissimilar. We then modified the community composition by turning the rewarding inflorescences into unrewarding and adding rewarding inflorescences of a novel flower colour. In the short term, bees trained to similar rather than dissimilar inflorescences experienced higher costs of foraging (decreased foraging speed and accuracy) in the novel community. The colonies differed in their speed-accuracy trade-off. In the longer term, bees adapted their foraging behaviour to the novel community composition by increasingly visiting the novel rewarding inflorescences.     

Getting to the start line: how bumblebees and honeybees are visually guided towards their first floral contact

Much of the literature on foraging behaviour in bees focuses on what they learn after they have had rewarded experience with flowers. This review focuses on how honeybees and bumblebees are drawn to candidate food sources in the first place: the foundation on which learning is built. Prior to rewarded foraging experience, flower-naïve bumblebees and honeybees rely heavily on visual cues to discover their first flower. This review lists methodological issues that surround the study of flower-naïve behaviour and describes technological advances. The role of distinct visual properties of flowers in attracting bees is considered: colour, floral size, patterning and social cues. The research reviewed is multi-disciplinary and takes the perspectives of both the bees and the plants they visit. Several avenues for future research are proposed.     

Buzz-pollination in Neotropical bees:genus-dependent frequencies and lack of optimal frequency for pollen release

Over 50 genera of bees release pollen from flower anthers using thoracic vibrations,a phenomenon known as buzz-pollination.The efficiency of this process is directly affected by the mechanical properties of the buzzes,namely the duration,amplitude,and frequency.Nonetheless,although the effects of the former two properties are well described,the role of buzz frequency on pollen release remains unclear.Furthermore,nearly all of the existing studies describing vibrational properties of natural buzz-pollination are limited to bumblebees(Bombus)and carpenter bees(Xvlocopa)constraining our current understanding of this behavior and its evolution.Therefore,we attempted to minimize this shortcoming by testing whether flower anthers exhibit optimal frequency for pollen release and whether bees tune their buzzes to match these(optimal)frequencies.If true,certain frequencies will trigger more pollen release and lighter bees will reach buzz frequencies closer to this optimum to compensate their smaller buzz amplitudes.Two strategies were used to test these hypotheses:(i)the use of(artificial)vibrational playbacks in a broad range of buzz frequencies and amplitudes to assess pollen release by tomato plants(Solarium Ivcopersicum L.)and(ii)the recording of natural buzzes of Neotropical bees visiting tomato plants during pollination.The playback experiment indicates that although buzz frequency does affect pollen release,no optimal frequency exists for that.In addition,the recorded results of natural buzz-pollination reveal that buzz frequencies vary with bee genera and are not correlated with body size.Therefore,neither bees nor plants are tuned to optimal pollen release frequencies.Bee frequency of buzz-pollination is a likely consequence of the insect flight machinery adapted to reach higher accelerations,while flower plant response to buzz-pollination is the likely result of its pollen granular properties.     

Colour preferences of flower-naive honeybees

Flower-naive honeybees Apis mellifera L. flying in an enclosure were tested for their colour preferences. Bees were rewarded once on an achromatic (grey, aluminium or hardboard), or on a chromatic (ultraviolet) disk. Since naive bees never alighted on colour stimuli alone, a scent was given in combination with colour. Their landings on twelve colour stimuli were recorded. Results after one reward (“first test”) were analysed separately from those obtained after few rewards (“late tests”).

1. After pre-training to achromatic signals, bees preferred, in the first test, bee-uv-blue and bee-green colours. With increasing experience, the original preference pattern persisted but the choice of bee-blue and bee-green colours increased.
2. Neither colour distance of the test stimuli to the background or to the pre-training signal, nor their intensity, nor their green contrast, accounted for the colour choice of bees. Choices reflected innate preferences and were only associated with stimulus hue.
3. Bees learned very quickly the pre-trained chromatic stimulus, the original colour preferences being thus erased.
4. Colour preferences were strongly correlated with flower colour and its associated nectar reward, as measured in 154 flower species.
5. Colour preferences also resemble the wavelength dependence of colour learning demonstrated in experienced bees.

     

Consequences of floral complexity for bumblebee-mediated geitonogamous self-pollination in Salvia nipponica Miq. (Labiatae)

Abstract.— I address how floral complexity influences geitonogamous self-pollination through manipulation of pollinator behavior in Salvia nipponica . The pivoting stamens of S. nipponica hinder nectar-collecting bumblebees from crawling into flowers, increasing the probing time per flower. I predicted that longer probing times would reduce the relative cost of moving between plants, causing bees to leave plants earlier. To test this prediction, I simplified S. nipponica flowers by removing the stamens from all open flowers within a 75-m² quadrat. Bumblebees probed these flowers more quickly than intact flowers, but the stamen removal affected neither the frequency of flower revisitation nor the flight distance between plants. In response to the decrease in the probing time per flower, bees probed more flowers on these plants. Therefore, in S. nipponica , floral complexity reduces the opportunity for geitonogamous self-pollination. Stamen removal also increased bee visitation per flower, suggesting that this sort of complexity deters visitation. To keep complex flowers attractive, therefore, selection might increase floral rewards or longevity. Floral complexity might evolve in an integrative manner with the rest of the floral phenotype.     

Pollination biology of the sclerophyllous shrub Pultenaea villosa Willd. (Fabaceae) in southeast Queensland, Australia

The pollination biology of the common shrub Pultenaea villosa Willd. was examined in a subtropical dry sclerophyll forest in eastern Australia. We determined floral phenology and morphology, the timing of stigma receptivity and anther dehiscence, nectar availability, the plant breeding system, and flower visitors. The shrub's flowers are typical zygomorphic pea flowers with hidden floral rewards and reproductive structures. These flowers require special manipulation for insect access. A range of insects visited the flowers, although bees are predicted to be the principle pollinators based on their frequency on the flowers and their exclusive ability to operate the wing and keel petals to access the reproductive structures. Nectar and pollen are offered as rewards and were actively collected by bees. Nectar is offered to visitors in minute amounts at the base of the corolla. In Toohey Forest, P. villosa flowers in spring and is the most abundant floral resource in the understory of the forest at this time. The breeding system experiment revealed that P. villosa requires outcrossing for high levels of seed set and that the overlap of stigma receptivity and pollen dehiscence within the flower suggests the potential for self-incompatibility.