Timing of invasive pollen deposition influences pollen tube growth and seed set in a native plant

Invasive plants may threaten the reproductive success of native sympatric plants by modifying the pollination process. One potential mechanism takes place through the deposition of invasive pollen onto native stigmas when pollinators are shared among species. We explore how pollen from the invasive plant Brassica nigra influences pre- and post-fertilization stages in the native plant Phacelia parryi, through a series of hand pollination experiments. These two species share pollinators to a high degree. P. parryi flowers were hand-pollinated with either pure conspecific pollen (the control) or with B. nigra pollen applied prior to, simultaneously with, or following conspecific pollen. Application of B. nigra pollen lowered seed set, with the simultaneous application resulting in the highest reduction. Pollen tube growth was also influenced by the presence of invasive pollen, with fewer conspecific pollen tubes reaching the base of P. parryi styles in treatments where B. nigra pollen was applied prior to or simultaneously with conspecific pollen. The deleterious effects of invasive pollen on native seed set in this study are likely not due to loss of stigmatic receptivity since seed set was less affected when heterospecific pollen was applied prior to conspecific pollen, but may instead involve interactions between interspecific pollen grains on the stigma or within the style. Our study highlights the importance of timing of foreign pollen deposition on native stigmas and suggests that interspecific pollen transfer between native and exotic plants may be an important mechanism of competition for pollination in invaded plant communities.     

Increased relative abundance of an invasive competitor for pollination, Lythrum salicaria, reduces seed number in Mimulus ringens

When exotic plant species share pollinators with native species, competition for pollination may lower the reproductive success of natives by reducing the frequency and/or quality of visits they receive. Exotic species often become numerically dominant in plant communities, and the relative abundance of these potential competitors for pollination may be an important determinant of their effects on the pollination and reproductive success of co-occurring native species. Our study experimentally tests whether the presence and abundance of an invasive exotic, Lythrum salicaria L. (Lythraceae), influences reproductive success of a co-flowering native species, Mimulus ringens L. (Phrymaceae). We also examine the mechanisms of competition for pollination and how they may be altered by changes in competitor abundance. We found that the presence of Lythrum salicaria lowered mean seed number in Mimulus ringens fruits. This effect was most pronounced when the invasive competitor was highly abundant, decreasing the number of seeds per fruit by 40% in 2006 and 33% in 2007. Reductions in the number of seeds per fruit were likely due to reduced visit quality resulting from Mimulus pollen loss when bees foraged on neighboring Lythrum plants. This study suggests that visit quality to natives may be influenced by the presence and abundance of invasive flowering plants.     

Relative abundance of an invasive alien plant affects native pollination processes

One major characteristic of invasive alien species is their occurrence at high abundances in their new habitat. Flowering invasive plant species that are visited by native insects and overlap with native plant species in their pollinators may facilitate or disrupt native flower visitation and fertilisation by forming large, dense populations with high numbers of flowers and copious rewards. We investigated the direction of such a proposed effect for the alien invasive Rhododendron ponticum in Irish habitats. Flower visitation, conspecific and alien pollen deposition, fruit and seed set were measured in a self-compatible native focal plant, Digitalis purpurea, and compared between field sites that contained different relative abundances of R. ponticum. Flower visitation was significantly lower at higher alien relative plant abundances than at lower abundances or in the absence of the alien. Native flowers experienced a significant decrease in conspecific pollen deposition with increasing alien abundance. Heterospecific pollen transfer was very low in all field sites but increased significantly with increasing relative R. ponticum abundance. However, lower flower visitation and lower conspecific pollen transfer did not alter reproductive success of D. purpurea. Our study shows that indirect interactions between alien and native plants for pollination can be modified by population characteristics (such as relative abundance) in a similar way as interactions among native plant species. In D. purpurea, only certain aspects of pollination and reproduction were affected by high alien abundances which is probably a result of high resilience due to a self-compatible breeding system. Native species that are more susceptible to pollen limitation are more likely to experience fitness disadvantages in habitats with high relative alien plant abundances.     

Plant–pollinator interactions between an invasive and native plant vary between sites with different flowering phenology

Floral displays of invasive plants have positive and negative impacts on native plant pollination. Invasive plants may also decrease irradiance, which can lead to reduced pollination of native plants. The effects of shade and flowers of invasive plant species on native plant pollination will depend on overlap in flowering phenologies. We examined the effect of the invasive shrub Lonicera maackii on female reproductive success of the native herb Hydrophyllum macrophyllum at two sites: one with asynchronous flowering phenologies (slight overlap) and one with synchronous (complete overlap). At each site, we measured light availability, pollinator visitation, pollen deposition, and seed set of potted H. macrophyllum in the presence and absence of L. maackii. At both sites, understory light levels were lower in plots containing L. maackii. At the asynchronous site, H. macrophyllum received fewer pollinator visits in the presence of L. maackii, suggesting shade from L. maackii reduced visitation to H. macrophyllum. Despite reduced visitation, H. macrophyllum seed set did not differ between treatments. At the synchronous site, H. macrophyllum received more pollinator visits and produced more seeds per flower in the presence of co-flowering L. maackii compared to plots in which L. maackii was absent, and conspecific pollen deposition was positively associated with seed set. Our results support the hypothesis that co-flowering L. maackii shrubs facilitated pollination of H. macrophyllum, thereby mitigating the negative impacts of shade, leading to increased seed production. Phenological overlap appears to influence pollinator-mediated interactions between invasive and native plants and may alter the direction of impact of L. maackii on native plant pollination.     

Shading by invasive shrub reduces seed production and pollinator services in a native herb

Plant invasions disrupt native plant reproduction directly via competition for light and other resources and indirectly via competition for pollination. Furthermore, shading by an invasive plant may reduce pollinator visitation and therefore reproduction in native plants. Our study quantifies and identifies mechanisms of these direct and indirect effects of an invasive shrub on pollination and reproductive success of a native herb. We measured pollinator visitation rate, pollen deposition, and female reproductive success in potted arrays of native Geranium maculatum in deciduous forest plots invaded by the non-native shrub Lonicera maackii and in two removal treatments: removal of aboveground L. maackii biomass and removal of flowers. We compared fruit and seed production between open-pollinated and pollen-supplemented plants to test for pollen and light limitation of reproduction. Plots with L. maackii had significantly lower light, pollinator visitation rate, and conspecific pollen deposition to G. maculatum than biomass removal plots. Lonicera maackii flower removal did not increase pollinator visitation or pollen deposition compared to unmanipulated invaded plots, refuting the hypothesis of competition for pollinators. Thus, pollinator-mediated impacts of invasive plants are not limited to periods of co-flowering or pollinator sharing between potential competitors. Geranium maculatum plants produced significantly fewer seeds in plots containing L. maackii than in plant removal plots. Seed set was similar between pollen-supplemented and open-pollinated plants, but pollen-supplemented plants exhibited higher seed set in plant removal plots compared to invaded plots. Therefore, we conclude that the mechanism of impact of L. maackii on G. maculatum reproduction was increased understory shade.     

Effects of multiple competitors for pollination on bumblebee foraging patterns and Mimulus ringens reproductive success

When co‐occurring plant species overlap in flowering phenology they may compete for the service of shared pollinators. Competition for pollination may lower plant reproductive success by reducing the number of pollinator probes or by decreasing the quality of pollen transport to or from a focal species. Pair‐wise interactions between plants sharing pollinators have been well documented. However, relatively few studies have examined interactions for pollination among three or more plant species, and little is known about how the outcomes and mechanisms of competition for pollination may vary with competitor species composition. To better understand how the dynamics of competition for pollination may be influenced by changes in the number of competitors, we manipulated the presence of two competitors, Lythrum salicaria and Lobelia siphilitica, and quantified reproductive success for a third species, Mimulus ringens. Patterns of pollinator preference and interspecific transitions in mixed‐species arrays were significantly influenced by the species composition of competitor plants present. Both pair‐wise and three‐species competition treatments led to a similar ～ 40% reduction in Mimulus ringens seed set. However, the patterns of pollinator foraging we observed suggest that the relative importance of different mechanisms of competition for pollination may vary with the identity and number of competitors present. This variation in mechanisms of competition for pollination may be especially important in diverse plant communities where many species interact through shared pollinators.     

Comparing the reproductive success and pollination biology of an invasive plant to its rare and common native congeners: a case study in the genus <Emphasis Type="Italic">Cirsium</Emphasis> (Asteraceae)

Previous studies have examined an association between reproductive success and pollination biology of rare versus widespread species through pair-wise comparisons of native and invasive congeners or rare and common congeners. To determine the importance of reproductive success and pollination biology for an invasive thistle, Cirsium vulgare, we compared it in its invaded range to five, co-occurring native Cirsium species that range from rare to common. Native study species include C. fontinale var. fontinale, C. andrewsii, C. brevistylum, C. occidentale, and C. quercetorum. We compared all species’ reproductive success, insect visitation rate and composition, autonomous self-pollination, and level of pollen limitation in multiple populations. Species differed in their reproductive success; the invasive C. vulgare produced more flower heads per plant than most native species. C. vulgare attracted more visitors than its congeners. In addition, reproductive success and insect visitation significantly varied between populations within species, mainly due to aphid infestation in one population of C. occidentale. Unlike the rare species (C. fontinale and andrewsii), C. vulgare did not require a pollinator for high-levels of seed production. The remaining native species set fewer seeds than C. vulgare without a pollinator. However, differences in insect visitation and autonomous self-pollination did not lead to differences in pollen limitation across species or between populations. This result suggests that factors other than pollination biology determine the difference in reproductive success of these species. However, high levels of autonomous self-pollination and generalist insect visitation may allow the invasive C. vulgare to easily establish new populations from low numbers of propagules. Our study provides one contrast that should build towards a larger comparative analysis to examine general patterns in the relationship between reproductive success, pollination biology, rare and invasive species, and our ability to predict biological invasions in introduced species.     

Effect of invader removal: pollinators stay but some native plants miss their new friend

Removal of invasive species often benefits biological diversity allowing ecosystems’ recovery. However, it is important to assess the functional roles that invaders may have established in their new areas to avoid unexpected results from species elimination. Invasive animal-pollinated plants may affect the plant–pollination interactions by changing pollinator availability and/or behaviour in the community. Thus, removal of an invasive plant may have important effects on pollinator community that may then be reflected positive or negatively on the reproductive success of native plants. The objective of this study was to assess the effect of removing Oxalis pes-caprae, an invasive weed widely spread in the Mediterranean basin, on plant–pollinator interactions and on the reproductive success of co-flowering native plants. For this, a disturbed area in central Portugal, where this species is highly abundant, was selected. Visitation rates, natural pollen loads, pollen tube growth and natural fruit set of native plants were compared in the presence of O. pes-caprae and after manual removal of their flowers. Our results showed a highly resilient pollination network but also revealed some facilitative effects of O. pes-caprae on the reproductive success of co-flowering native plants. Reproductive success of the native plants seems to depend not only on the number and diversity of floral visitors, but also on their efficiency as pollinators. The information provided on the effects of invasive species on the sexual reproductive success of natives is essential for adequate management of invaded areas.     

Interspecific pollen transfer as a mechanism of competition: Consequences of foreign pollen contamination for seed set in the alpine wildflower,Polemonium viscosum

Summary Earlier studies showed that under natural conditions foreign pollen receipt is correlated with loss of seed set in populations of P. viscosum. Here, we report on experimental pollinations using foreign pollen from two co-flowering species that share pollinators with P. viscosum. Experiments were designed to ascertain whether improper pollen transfer causes reduced seed set under controlled conditions and, if so, to determine the stage at which interference occurs. In flowers pre-treated by pollination with either Mertensia or Castilleja pollen, adherence of subsequently deposited conspecific pollen grains was unaffected. However, conspecific grains had significantly lower germination and ovule fertilization success in flowers receiving foreign pollen than in controls. These results suggest that pollinator infidelity in high alpine plant communities can reduce female reproductive success of P. viscosum by inhibiting the performance of conspecific pollen grains. Negative effects of Mertensia and Castilleja pollen were comparable despite large interspecific differences in pollen grain morphology. Accordingly, susceptibility to foreign pollen interference appears to be an intrinsic property of P. viscosum that is not dependent on the unique structural properties of pollen produced by competitive species.     

Heterospecific pollen deposition: does diversity alter the consequences?

? In natural communities, plants can receive pollen from multiple heterospecifics as well as conspecifics. However, studies on the effects of interspecific pollen transfer have focused on interactions between species pairs. The potential exists for diverse interactions among heterospecific pollen (HP) grains on the stigma, and for these to affect plant reproduction, alone or in combination with conspecific pollen (CP) loss, but these interactions have not yet been explored. ? We used hand-pollinations to simulate increasing community diversity and CP loss on Mimulus guttatus stigmas. We used pollen mixes of one to three heterospecific donors to determine how species composition and CP load size affect seed production and to characterize the mechanisms underlying fertilization failure. ? Heterospecific pollen deposition reduced M. guttatus seed production and while the effect increased with the number of heterospecific donors, the strength depended on species composition and was independent of conspecific load size. Different types of interactions (additive and synergistic) are hypothesized to underlie the diverse effects on M. guttatus reproductive success. ? Our results suggest that an increase in the diversity of heterospecific donors will not always lead to a greater decrease in fitness because multispecies effects depend on the interacting species.     

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.     

Reproductive interactions mediated by flowering overlap in a temperate hummingbird–plant assemblage

Pollinator‐mediated competition through shared pollinators can lead to segregated flowering phenologies, but empirical evidence for the process responsible for this flowering pattern is sparse. During two flowering seasons, we examined whether increasing overlap in flowering phenology decreased conspecific pollination, increased heterospecific pollination, and depressed seed output in the seven species composing a hummingbird–plant assemblage from the temperate forest of southern South America. Overall trends were summarized using meta‐analysis. Despite prevailing negative associations, relations between phenological overlap and conspecific pollen receipt varied extensively among species and between years. Heterospecific pollen receipt was low and presumably of limited biological significance. However, our results supported the hypothesis that concurrent flowering promotes interspecific pollen transfer, after accounting for changes in the abundance of conspecific flowers. Seed output was consistently reduced during maximum phenological overlap during the first flowering season because of limited fruit set. Responses varied more during the second year, despite an overall negative trend among species. Relations between estimated effects of phenological overlap on pollination and seed output, however, provided mixed evidence that conspecific pollen loss during pollinator visits to foreign flowers increases pollen limitation. By flowering together, different plant species might benefit each other's pollination by increasing hummingbird recruitment at the landscape level. Nevertheless, our results are mostly consistent with the hypothesis of pollinator‐mediated competition shaping the segregated flowering pattern reported previously for this temperate plant assemblage. The mechanisms likely involve effects on male function, whereby pollen‐transport loss during heterospecific flower visits limit pollen export, and more variable effects on female function through pollen limitation.     

Effects of conspecific and heterospecific floral density on the pollination of two related rewarding orchids

Variation in within-population floral density can affect interactions between plants and pollinators, resulting in variable pollen export for plants. We investigated the effects of conspecific and heterospecific floral densities on pollination success both of two related, self-compatible, nectar-rewarding orchid species in Ireland, Spiranthes romanzoffiana (rare and listed as endangered) and its congener, S. spiralis (more abundant and not of conservation concern). Floral densities, insect visitation rates, and orchid pollen transport were recorded in multiple quadrats in four populations of both orchid species over their flowering season. We found that conspecific and heterospecific co-flowering plant density affected pollination in both orchid species. For S. romanzoffiana, higher heterospecific density increased pollen removal. For S. spiralis, higher conspecific visitation increased pollen removal and increased heterospecific density decreased pollen deposition. In addition, increased conspecific density increased pollen deposition in both species. This study shows that plants may interact to facilitate or compete for different components of the pollination process, namely; pollinator attraction, pollen removal and deposition. Such interactions have immediate consequences for endangered plant species, as increases in both conspecific and heterospecific coflowering density may ameliorate the negative effects of rarity on pollination, hence overall reproductive success.     

Outcrossing breeding system does not compromise invasiveness in Buddleja davidii

Introductions of alien plant species are often likely to consist of a few individuals. Thus, invasion success may strongly depend on their reproductive biology. A high number of self-compatible plants species are known to be successful colonizers of new habitats, even able to establish populations from single propagules. However, many other invasive species require pollen vectors. Here, we investigated the mating system of Buddleja davidii, a fast growing shrub native to China that colonizes quickly in disturbed habitats such as quarries, river banks, along railways and roads, both in its native and invasive regions. It was intentionally introduced to Europe as an ornamental plant because of its fragrant and showy flowers. We additionally studied its vulnerability to biparental inbreeding depression by performing a controlled crossing experiment using pollen from the same population or from geographically close and distant populations, respectively. As a measure for pollination success, we used capsule weight, seed number per capsule and seed weight for each treatment.     

The presence of the invasive plant Solanum elaeagnifolium deters honeybees and increases pollen limitation in the native co-flowering species Glaucium flavum

Invasive plants can impact biodiversity and ecosystem functioning by displacing native plants and crop species due to competition for space, nutrients, water and light. The presence of co-flowering invasives has also been shown to affect some native plants through the reduction in pollinator visitation or through the deposition of heterospecific pollen on the native’s stigmas leading to stigma clogging. We examined the impact of the invasive plant Solanum elaeagnifolium Cavanilles (silver-leafed nightshade), native to South and Central America and South-western parts of North America, on the seed set of the native Glaucium flavum Crantz (yellow-horned poppy) on Lesvos Island, Greece. To do this we measured seed set and visitation rates to G. flavum before and after the placement of potted individuals of the invasive near the native plants. In addition, we hand-crossed G. flavum flowers with super-optimal amounts of conspecific pollen, bagged flowers to measure the rate of spontaneous selfing, and applied self-pollen to measure self-compatibility of G. flavum. The hand-selfing treatment resulted in very low seed set, which indicates that G. flavum is to a large degree self-incompatible and highlights the plant’s need for insect-mediated outcrossing. We show that the presence of the invasive significantly enhanced pollen limitation, although the overall visitation rates were not reduced and that this increase is due to a reduction in honeybee visitation in the presence of the invasive resulting in reduced pollination.     

Prezygotic barriers to gene flow between <Emphasis Type="Italic">Taraxacum ceratophorum</Emphasis> and the invasive <Emphasis Type="Italic">Taraxacum officinale</Emphasis> (Asteraceae)

Prezygotic reproductive barriers limit interspecific gene flow between congeners. Here, I examine the strength of floral isolation and interspecific pollen-pistil barriers between an invasive apomictic, Taraxacum officinale, and the indigenous sexual alpine dandelion, Taraxacum ceratophorum. Experimental arrays of either native inflorescences or a mixture of native and exotic inflorescences were used to examine insect preference and to track movement of a pollen analog. Using hand-pollinations, conspecific and heterospecific pollen germination success on native stigmas was compared. To additionally test for interspecific pollen competition, T. ceratophorum plants received one of three possible hand-pollinations: control conspecific pollination, concomitant conspecific and heterospecific pollination (mixed), or conspecific pollen followed by heterospecific pollen 15 min later (staggered). Floral isolation was negligible as no insect preference was detected. On a presence/absence basis, florets on native inflorescences received slightly less pollen analog from heterospecific donors than from conspecific donors; however, the amount of dye particles transferred from either Taraxacum species to stigmas on recipient T. ceratophorum inflorescences was equivalent. In contrast to weak floral isolation, strong pollen germination and pollen competition barriers should reduce the potential for hybridization. Heterospecific T. officinale pollen exhibited reduced germination success on T. ceratophorum stigmas in comparison to conspecific pollen. Furthermore, a significant pollen-competition effect on the percentage of hybrid offspring was detected only when T. officinale preceded T. ceratophorum pollen by 15 min. This result indicates that conspecific pollen out-competes heterospecific pollen but further suggests that biotic and abiotic factors reducing pollen accrual rates may partially remove barriers to natural hybridization.     

Multiple pollinator visits to Mimulus ringens (Phrymaceae) flowers increase mate number and seed set within fruits

The timing and effectiveness of pollinator visitation to flowers is an important factor influencing mating patterns and reproductive success. Multiple pollinator probes to a flower may increase both the quantity and genetic diversity of progeny, especially if single probes deposit insufficient pollen for maximal seed set or if the interval between probes is brief. When pollen carryover is limited, sequential pollen loads may also differ markedly in sire representation. We hypothesized that these conditions help explain high levels of multiple paternity in Mimulus ringens fruits. We documented all bee visits to individual flowers, quantified resulting seed set, and determined paternity for 20 seeds per fruit. Most (76%) flowers received multiple probes, and the interval between probes was usually <30 min. Flowers probed multiple times produced 44% more seeds than flowers probed once. All fruits were multiply sired. Flowers receiving a single probe averaged 3.12 outcross sires per fruit, indicating that single probes deposit pollen from several donors. Multiple paternity was even greater after three or more probes (4.92 outcross sires), demonstrating that sequential visits bring pollen from donors not represented in the initial probe.     

Experimental reduction of pollinator visitation modifies plant–plant interactions for pollination

The strength of interactions between plants for pollination depends on the abundance of plants and pollinators in the community. The abundance of pollinators may influence plant associations and densities at which individual fitness is maximized. Reduced pollinator visitation may therefore affect the way plant species interact for pollination. We experimentally reduced pollinator visitation to six pollinator‐dependent species (three from an alpine and three from a lowland community in Norway) to study how interactions for pollination were modified by reduced pollinator availability. We related flower visitation, pollen limitation and seed set to density of conspecifics and pollinator‐sharing heterospecifics inside 30 dome‐shaped cages partially covered with fishnet (experimental plots) and in 30 control plots. We expected to find stronger interactions between plants in experimental compared to controls plots. The experiment modified plant–plant interactions for pollination in all the six species; although for two of them neighbourhood interactions did not affect seed set. The pollen limitation and seed set data showed that reduction of pollinator visits most frequently resulted in novel and/or stronger interactions between plants in the experimental plots that did not occur in the controls. Although the responses were species‐specific, there was a tendency for increasing facilitative interactions with conspecific neighbours in experimental plots where pollinator availability was reduced. Heterospecifics only influenced pollination and fecundity in species from the alpine community and in the experimental plots, where they competed with the focal species for pollination. The patterns observed for visitation rates differed from those for fecundity, with more significant interactions between plants in the controls in both communities. This study warns against the exclusive use of visitation data to interpret plant–plant interactions for pollination, and helps to understand how plant aggregations may buffer or intensify the effects of a pollinator loss on plant fitness.     

Asymmetrical conspecific seed-siring advantage between Silene latifolia and S. dioica

Background and Aims

Silene dioica and S. latifolia experience only limited introgression despite overlapping flowering phenologies, geographical distributions, and some pollinator sharing. Conspecific pollen precedence and other reproductive barriers operating between pollination and seed germination may limit hybridization. This study investigates whether barriers at this stage contribute to reproductive isolation between these species and, if so, which mechanisms are responsible.

Methods

Pollen-tube lengths for pollen of both species in styles of both species were compared. Additionally, both species were pollinated with majority S. latifolia and majority S. dioica pollen mixes; then seed set, seed germination rates and hybridity of the resulting seedlings were determined using species-specific molecular markers.

Key Results

The longest pollen tubes were significantly longer for conspecific than heterospecific pollen in both species, indicating conspecific pollen precedence. Seed set but not seed germination was lower for flowers pollinated with pure heterospecific versus pure conspecific pollen. Mixed-species pollinations resulted in disproportionately high representation of nonhybrid offspring for pollinations of S. latifolia but not S. dioica flowers.

Conclusions

The finding of conspecific pollen precedence for pollen-tube growth but not seed siring in S. dioica flowers may be explained by variation in pollen-tube growth rates, either at different locations in the style or between leading and trailing pollen tubes. Additionally, this study finds a barrier to hybridization operating between pollination and seed germination against S. dioica but not S. latifolia pollen. The results are consistent with the underlying cause of this barrier being attrition of S. dioica pollen tubes or reduced success of heterospecifically fertilized ovules, rather than time-variant mechanisms. Post-pollination, pre-germination barriers to hybridization thus play a partial role in limiting introgression between these species.     

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.