Relationships between the floral neighborhood and individual pollen limitation in two self-incompatible herbs

Local flower density can affect pollen limitation and plant reproductive success through changes in pollinator visitation and availability of compatible pollen. Many studies have investigated the relationship between conspecific density and pollen limitation among populations, but less is known about within-population relationships and the effect of heterospecific flower density. In addition, few studies have explicitly assessed how the spatial scales at which flowers are monitored affect relationships. We investigated the effect of floral neighborhood on pollen limitation at four spatial scales in the self-incompatible herbs Armeria maritima spp. maritima and Ranunculus acris spp. acris. Moreover, we measured pollen deposition in Armeria and pollinator visits to Ranunculus. There was substantial variation in pollen limitation among Armeria individuals, and 25% of this variation was explained by the density of compatible and heterospecific flowers within a 3 m circle. Deposition of compatible pollen was affected by the density of compatible and incompatible inflorescences within a 0.5 m circle, and deposition of heterospecific pollen was affected by the density of heterospecific flowers within a 2 m circle. In Ranunculus, the number of pollinator visits was affected by both conspecific and heterospecific flower densities. This did not, however, result in effects of the floral neighborhood on pollen limitation, probably due to an absence of pollen limitation at the population level. Our study shows that considerable variation in pollen limitation may occur among individuals of a population, and that this variation is partly explained by floral neighborhood density. Such individual-based measures provide an important link between pollen limitation theory, which predicts ecological and evolutionary causes and consequences for individual plants, and studies of the effects of landscape fragmentation on plant species persistence. Our study also highlights the importance of considering multiple spatial scales to understand the spatial extent of pollination processes within a population.     

Do local conspecific density and floral display size influence fruit set via pollinator visitation in Orchis militaris?

Plant density varies naturally, from isolated plants to clumped individuals, and this can influence pollinator foraging behaviour and plant reproductive success. In addition, the effect of conspecific density on reproduction may depend on the pollination system, and deceptive species differ from rewarding ones in this regard, a high density being often associated with low fruit set in deceptive plants. In our study, we aimed to determine how local conspecific density and floral display size (i.e. number of flowers per plant) affect fruit set in a deceptive orchid (Orchis militaris) through changes in pollinator visitation. We measured fruit set in a natural population and recorded pollinator abundance and foraging behaviour within plots of different O. militaris densities. Detailed data were recorded for the most abundant potential pollinators of O. militaris, i.e. solitary bees. Floral display size was negatively correlated to fruit set in medium‐density plots, but uncorrelated in low‐ and high‐density plots. Plot density had no effect on solitary bee abundance and visitation, which may be due to low pollinator abundance within the study site. The proportion of visited flowers per inflorescence was negatively influenced by floral display size, which is in line with previous studies. In addition, solitary bees spent decreasing time in successive flowers within an inflorescence, and the time spent per flower was negatively affected by ambient temperature. Our results suggest that pollinator behaviour during visitation is poorly linked to pollen deposition and reproductive success in O. militaris.     

Effects of floral restrictiveness and stigma size on heterospecific pollen receipt in a prairie community

Plant species vary greatly in the degree to which floral morphology restricts access to the flower interior. Restrictiveness of flower corollas may influence heterospecific pollen receipt, but the impact of floral morphology on heterospecific pollen transfer has received little attention. We characterized patterns of pollinator visitation and quantities of conspecific and heterospecific pollen receipt for 29 species with a range of floral morphologies in a prairie community dominated by the introduced plant Euphorbia esula (leafy spurge) which has an unrestrictive morphology. Pollinator overlap was significantly greater between Euphorbia and other unrestrictive flowers than restrictive flowers. Compared to flowers with restrictive morphologies, unrestrictive flowers received significantly more Euphorbia pollen, more heterospecific pollen from other sources, and a greater diversity of pollen species, but not more conspecific pollen. However, stigmatic surface area was significantly larger for flowers with unrestrictive morphologies, and the density of Euphorbia and other heterospecific pollen per stigmatic area did not significantly differ between flower types. These findings suggest that the smaller stigma size in restrictive flowers partly accounts for their decreased heterospecific pollen receipt, but that restrictiveness also allows species to increase the purity of pollen loads they receive. Given that restrictive flowers receive fewer heterospecific pollen grains but at a higher density, the effect of restrictiveness on fecundity depends on whether absolute quantity or density of heterospecific pollen affects fecundity more. Our results also indicate that abundant neighbors are not necessarily important heterospecific pollen sources since Euphorbia pollen was rarely abundant on heterospecifics.     

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.     

Effects of distance from models on the fitness of floral mimics

• Rewardless plants can attract pollinators by mimicking floral traits of rewarding heterospecific plants. This should result in the pollination success of floral mimics being dependent on the relative abundance of their models, as pollinator abundance and conditioning on model signals should be higher in the vicinity of the models. However, the attraction of pollinators to signals of the models may be partially innate, such that spatial isolation of mimics from model species may not strongly affect pollination success of mimics.
• We tested whether pollination rates and fruit set of the rewardless orchid Disa pulchra were influenced by proximity and abundance of its rewarding model species, Watsonia lepida.
• Pollination success of the orchid increased with proximity to the model species, while fruit set of the orchid increased with local abundance of the model species. Orchids that were experimentally translocated outside the model population experienced reduced pollinaria removal and increased pollinator‐mediated self‐pollination.
• These results confirm predictions that the pollination success of floral mimics should be dependent on the proximity and abundance of model taxa, and thus highlight the importance of ecological facilitation among species involved in mimicry systems.

     

Pollen competition between two sympatric Orchis species (Orchidaceae): the overtaking of conspecific of heterospecific pollen as a reproductive barrier

The frequency of hybrid formation in angiosperms depends on how and when heterospecific pollen is transferred to the stigma, and on the success of that heterospecific pollen at fertilising ovules. We applied pollen mixtures to stigmas to determine how pollen interactions affect siring success and the frequency of hybrid formation between two species of Mediterranean deceptive orchid. Plants of Orchis italica and O. anthropophora were pollinated with conspecific and heterospecific pollen (first conspecific pollen then heterospecific pollen and vice versa) and molecular analysis was used to check the paternity of the seeds produced. In this pair of Mediterranean orchids, competition between conspecific and heterospecific pollen functions as a post‐pollination pre‐zygotic barrier limiting the frequency of the formation of hybrids in nature. Flowers pollinated with heterospecific pollen can remain receptive for the arrival of conspecific pollen for a long time. There is always an advantage of conspecific pollen for fruit formation, whether it comes before or after heterospecific pollen, because it overtakes the heterospecific pollen. The conspecific pollen advantage exhibited in O. italica and O. anthropophora is likely to result from the reduced germination of heterospecific pollen or retarded growth of heterospecific pollen tubes in the stigma and ovary. Overall, the results indicate that our hybrid zone represents a phenomenon of little evolutionary consequence, and the conspecific pollen advantage maintains the genetic integrity of the parental species.     

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

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

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.     

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.     

Disentangling multiple drivers of pollination in a landscape-scale experiment

Animal pollination is essential for the reproductive success of many wild and crop plants. Loss and isolation of (semi-)natural habitats in agricultural landscapes can cause declines of plants and pollinators and endanger pollination services. We investigated the independent effects of these drivers on pollination of young cherry trees in a landscape-scale experiment. We included (i) isolation of study trees from other cherry trees (up to 350 m), (ii) the amount of cherry trees in the landscape, (iii) the isolation from other woody habitats (up to 200 m) and (iv) the amount of woody habitats providing nesting and floral resources for pollinators. At the local scale, we considered effects of (v) cherry flower density and (vi) heterospecific flower density. Pollinators visited flowers more often in landscapes with high amount of woody habitat and at sites with lower isolation from the next cherry tree. Fruit set was reduced by isolation from the next cherry tree and by a high local density of heterospecific flowers but did not directly depend on pollinator visitation. These results reveal the importance of considering the plant''s need for conspecific pollen and its pollen competition with co-flowering species rather than focusing only on pollinators’ habitat requirements and flower visitation. It proved to be important to disentangle habitat isolation from habitat loss, local from landscape-scale effects, and direct effects of pollen availability on fruit set from indirect effects via pollinator visitation to understand the delivery of an agriculturally important ecosystem service.     

Congruence between visitation and pollen-transport networks in a California plant–pollinator community

Most recent studies describing pollination networks are based on observed flower visits, and few have explicitly tested if the floral visitors actually carry pollen. Since floral visitors can vary in their ability to remove and transfer pollen, it is important to show that visitation patterns reflect effective pollination. Given the difficulty of measuring per-visit pollen deposition at the community scale, a first step is to examine the amount of conspecific pollen carried by insect visitors. Here I compared the plant–animal visitation network with the pollen-transport network, estimated from insect pollen loads, for a montane meadow community from southern California, USA. Visitation and pollen-transport networks were positively associated with each other in both 2001 and 2002. However, the exclusion of visitors that do not carry any conspecific pollen reveals that pollen-transport networks are more specialized from the plants' perspective and that species are involved in fewer mutualistic interactions compared with estimates derived from visitation frequencies. Although conspecific pollen loads were smaller in 2002, bees tended to carry the largest conspecific loads in both years and were responsible for transporting the most pollen. These results suggest that, although visitation networks are suitable first-order approximations of pollination networks, information on which visitors carry conspecific pollen, and in what amounts, is crucial for distinguishing between antagonistic and mutualistic interactions.     

Mating system and the hybridization between self-compatible Phlox cuspidata and self-incompatible Phlox drummondii

Mating system can impact the frequency of hybridization and therefore the maintenance of species diversity. I evaluate the effects of weak self-incompatibility (SI) in Phlox cuspidata and SI in Phlox drummondii on mating success within species and on hybridization dynamics between species under controlled conditions. The effects of SI on hybridization frequency were assessed by manipulating the relatedness of conspecific pollen and the relative timing of pollen deposition in mixed-donor interspecific pollinations. Selfing as opposed to outcrossing increased hybridization by 16% in P. cuspidata maternal plants and by 48% in P. drummondii maternal plants because self pollen did not compete as well against heterospecific pollen. The relative timing of conspecific versus heterospecific pollen deposition also impacted hybridization. In self-compatible P. cuspidata, the deposition of self pollen 5 h earlier than heterospecific pollen decreased hybridization by 28%. In self-incompatible P. drummondii, a 5 h delay in the deposition of compatible conspecific pollen increased hybridization by 32%. In this hybrid system, early self-pollination can decrease hybridization (but increase inbreeding) by P. cuspidata maternal plants, and SI may increase hybridization by P. drummondii maternal plants.     

Unique growth paths of heterospecific pollen tubes result in late entry into ovules in the gynoecium of Sagittaria (Alismataceae)

• Pollen‐pistil interactions are a fundamental process in the reproductive biology of angiosperms and play a particularly important role in maintaining incipient species that exist in sympatry. However, the majority of previous studies have focused on species with syncarpous gynoecia (fused carpels) and not those with apocarpous gynoecia (unfused carpels).
• In the present study, we investigated the growth of conspecific pollen tubes compared to heterospecific pollen tubes in Sagittaria species, which have apocarpous gynoecia. We conducted controlled pollinations between S. pygmaea and S. trifolia and observed the growth of conspecific and heterospecific pollen tubes under a fluorescence microscope.
• Heterospecific and conspecific pollen tubes arrived at locules within the ovaries near simultaneously. However, conspecific pollen tubes entered into the ovules directly, whereas heterospecific tubes passed through the carpel base and adjacent receptacle tissue, to ultimately fertilize other unfertilized ovules. This longer route taken by heterospecific pollen tubes therefore caused a delay in the time required to enter into the ovules. Furthermore, heterospecific pollen tubes displayed similar growth patterns at early and peak pollination. The growth pattern of heterospecific pollen tubes at late pollination was similar to that of conspecific pollen tubes at peak pollination.
• Heterospecific and conspecific pollen tubes took different routes to fertilize ovules. A delayed entry of heterospecific pollen into ovules may be a novel mechanism of conspecific pollen advantage (CPA) for apocarpous species.

     

Floral density and co-occurring congeners alter patterns of selection in annual plant communities*

Although the evolution and diversification of flowers is often attributed to pollinator-mediated selection, interactions between co-occurring plant species can alter patterns of selection mediated by pollinators and other agents. The extent to which both floral density and congeneric species richness affect patterns of net and pollinator-mediated selection on multiple co-occurring species in a community is unknown and is likely to depend on whether co-occurring plants experience competition or facilitation for reproduction. We conducted an observational study of selection on four species of Clarkia (Onagraceae) and tested for pollinator-mediated selection on two Clarkia species in communities differing in congeneric species richness and local floral density. When selection varied with community context, selection was generally stronger in communities with fewer species, where local conspecific floral density was higher, and where local heterospecific floral density was lower. These patterns suggest that intraspecific competition at high densities and interspecific competition at low densities may affect the evolution of floral traits. However, selection on floral traits was not pollinator mediated in Clarkia cylindrica or Clarkia xantiana, despite variation in pollinator visitation and the extent of pollen limitation across communities for C. cylindrica. As such, interactions between co-occurring species may alter patterns of selection mediated by abiotic agents of selection.     

Pollination outcomes reveal negative density‐dependence coupled with interspecific facilitation among plants

Pollination is thought to be under positive density‐dependence, destabilising plant coexistence by conferring fitness disadvantages to rare species. Such disadvantage is exacerbated by interspecific competition but can be mitigated by facilitation and intraspecific competition. However, pollinator scarcity should enhance intraspecific plant competition and impose disadvantage on common over rare species (negative density‐dependence, NDD). We assessed pollination proxies (visitation rate, pollen receipt, pollen tubes) in a generalised plant community and related them to conspecific and heterospecific density, expecting NDD and interspecific facilitation due to the natural pollinator scarcity. Contrary to usual expectations, all proxies indicated strong intraspecific competition for common plants. Moreover interspecific facilitation prevailed and was stronger for rare than for common plants. Both NDD and interspecific facilitation were modulated by specialisation, floral display and pollinator group. The combination of intraspecific competition and interspecific facilitation fosters plant coexistence, suggesting that pollination can be a niche axis maintaining plant diversity.     

Interspecific pollinator movements reduce pollen deposition and seed production in Mimulus ringens (Phrymaceae)

Movement of pollinators between coflowering plant species may influence conspecific pollen deposition and seed set. Interspecific pollinator movements between native and showy invasive plants may be particularly detrimental to the pollination and reproductive success of native species. We explored the effects of invasive Lythrum salicaria on the reproductive success of Mimulus ringens, a wetland plant native to eastern North America. Pollinator flights between these species significantly reduced the amount of conspecific pollen deposited on Mimulus stigmas and the number of seeds in Mimulus fruits, suggesting that pollen loss is an important mechanism of competition for pollination. Although pollen loss is often attributed to pollen wastage on heterospecific floral structures, our novel findings suggest that grooming by bees as they forage on a competitor may also significantly reduce outcross pollen export and seed set in Mimulus ringens.     

Bumblebee visitation and seedset in Melampyrum pratense and Viscaria vulgaris: heterospecific pollen and pollen limitation

Summary Fruiting and seed set in two bumblebee-pollinated herbs, Melampyrum pratense L. (annual, Scrophulariaceae) and Viscaria vulgaris Bernh. (perennial, Caryophyllaceae) were studied on a dry meadow in southwestern Sweden in June 1986 and 1988. Both species produced seeds by self-fertilization. In Melampyrum (homogamous) fruiting and seed set by selfing were much lower than by natural pollination; in Viscaria (protandrous) fruiting by selfing and by natural polination were similar, but seed set per flower was lower by selfing than by natural pollination. Sequential hand pollinations increased seed set in Melampyrum, but not in Viscaria. Thus, the number of pollinations is important for high seed set in Melampyrum, and number of pollen grains deposited one pollination is important for high seed set in Viscaria. Late flowering resulted in the production of fewer seeds in both species, although the visitation rate in pure Viscaria stands was sufficient, because of limited resources. Pollen was the limiting resource in Viscaria, because hand pollination increased natural seed set. In Melampyrum pollen was limiting in 1988 but so were consumable resources, because the seedset decreased with time despite hand pollination. Pure stands of Viscaria had sced set similar to plants in mixed stands (with Melampyrum and Rhinanthus), although plants in mixed stands received fewer visits. Many seeds produced late in the season are the result of self pollination; emasculated Viscaria flowers had a very low seedset late in the season. Pollen loads containing approximately 50% heterospecific grains did not affect seed set in either species. Application of heterospecific (Lupinus) pollen to receptive Viscaria styles 6 h before conspecific pollen did not affect seed set.     

Interactions between a pollinating seed predator and its host plant: the role of environmental context within a population

Plant–insect interactions often are important for plant reproduction, but the outcome of these interactions may vary with environmental context. Pollinating seed predators have positive and negative effects on host plant reproduction, and the interaction outcome is predicted to vary with density or abundance of the partners. We studied the interaction between Silene stellata, an herbaceous perennial, and Hadena ectypa, its specialized pollinating seed predator. Silene stellata is only facultatively dependent upon H. ectypa for pollination because other nocturnal moth co‐pollinators are equally effective at pollen transfer. We hypothesized that for plants without conspecific neighbors, H. ectypa would have higher visitation rates compared to co‐pollinators, and the plants would experience lower levels of H. ectypa pollen deposition. We predicted similar oviposition throughout the study site but greater H. ectypa predation in the area without conspecific neighbors compared to plants embedded in a naturally high density area. We found that H. ectypa had consistently higher visitation than moth co‐pollinators in all host plant contexts. However, H. ectypa pollinator importance declined in areas with low conspecific density because of reduced pollen deposition, resulting in lower seed set. Conversely, oviposition was similar across the study site independent of host plant density. Greater likelihood of very high fruit predation combined with lower pollination by H. ectypa resulted in reduced S. stellata female reproductive success in areas with low conspecific density. Our results demonstrate local context dependency of the outcomes of pollinating seed predator interactions with conspecific host plant density within a population.     

POLLINATOR‐MEDIATED REPRODUCTIVE ISOLATION AMONG DIOECIOUS FIG SPECIES (FICUS,MORACEAE)

The extent of isolation among closely related sympatric plant species engaged in obligate pollination mutualisms depends on the fitness consequences of interspecies floral visitation. In figs (Ficus), interspecific gene flow may occur when pollinating wasps (Agaonidae) visit species other than their natal fig species. We studied reproductive isolation in a clade of six sympatric dioecious fig species in New Guinea. Microsatellite genotyping and Bayesian clustering analysis of the fig community indicated strong reproductive barriers among sympatric species. A total of 1–2% of fig populations consisted of hybrid individuals. A new experimental method of manipulating fig wasps investigated the reproductive consequences of conspecific and heterospecific pollinator visitation for both mutualists. Fig wasps introduced to Ficus hispidioides pollinated and oviposited in receptive figs. Seed development and seedling growth were largely comparable between conspecific and heterospecific crosses. Heterospecific pollinator fitness, however, was significantly less than that of conspecific pollinators. Heterospecific pollinators induced gall formation but offspring did not develop to maturity in the new host. Selection on pollinators maintaining host specificity appears to be an important mechanism of contemporary reproductive isolation among these taxa that could potentially influence their diversification.     

Incomplete partitioning of pollinators by Linum suffruticosum and its coflowering congeners

Premise

Linum suffruticosum shows variations in pollinator fit, pollen pickup, and local pollinators that predict pollen deposition rates. The species often coflowers with other Linum species using the same pollinators. We investigated whether L. suffruticosum trait variation could be explained by local patterns of pollinator sharing and associated evolution to reduce interspecific pollen transfer.

Methods

Pollinator observations were made in different localities (single species, coflowering with other congeners). Floral traits were measured to detect differences across populations and from coflowering species. Reproductive costs were quantified using interspecific hand pollinations and measures of pollen-tube formation, combined with observations of pollen arrival on stigmas and pollen-tube formation after natural pollination in allopatric and sympatric localities.

Results

The size and identity of the most important pollinator of L. suffruticosum and whether there was pollinator sharing with coflowering species appeared to explain floral trait variation related to pollinator fit. The morphological overlap of the flowers of L. suffruticosum with those of coflowering species varied, depending on coflowering species identity. A post-pollination incompatibility system maintains reproductive isolation, but conspecific pollen-tube formation was lower after heterospecific pollination. Under natural pollination at sites of coflowering with congeners, conspecific pollen-tube formation was lower than at single-species localities.

Conclusions

Trait variation in L. suffruticosum appears to respond to the most important local pollinator. Locally, incomplete pollinator partitioning might cause interspecific pollination, imposing reproductive costs. These reproductive costs may generate selection on floral traits for reduced morphological overlap with coflowering congeners, leading to the evolution of pollination ecotypes.