Pollinator identity and spatial isolation influence multiple paternity in an annual plant

The occurrence and extent of multiple paternity is an important component of variation in plant mating dynamics. However, links between pollinator activity and multiple paternity are generally lacking, especially for plant species that attract functionally diverse floral visitors. In this study, we separated the influence of two functionally distinct floral visitors (hawkmoths and solitary bees) and characterized their impacts on multiple paternity in a self‐incompatible, annual forb, Oenothera harringtonii (Onagraceae). We also situated pollinator‐mediated effects in a spatial context by linking variation in multiple paternity to variation in plant spatial isolation. We documented pronounced differences in the number of paternal sires as function of pollinator identity: on average, the primary pollinator (hawkmoths) facilitated mating with nearly twice as many pollen donors relative to the secondary pollinator (solitary bees). This effect was consistent for both isolated and nonisolated individuals, but spatial isolation imposed pronounced reductions on multiple paternity regardless of pollinator identity. Considering that pollinator abundance and pollen dispersal distance did not vary significantly with pollinator identity, we attribute variation in realized mating dynamics primarily to differences in pollinator morphology and behaviour as opposed to pollinator abundance or mating incompatibility arising from underlying spatial genetic structure. Our findings demonstrate that functionally distinct pollinators can have strongly divergent effects on polyandry in plants and further suggest that both pollinator identity and spatial heterogeneity have important roles in plant mating dynamics.     

Multiple paternity in fruits of Ipomopsis aggregata (Polemoniaceae)

Two different mechanisms can result in multiple paternity within fruits: deposition of a mixed pollen load due to carryover of pollen from flower to flower and multiple pollinator visits in close succession. I investigated the extent of multiple paternity within fruits of Ipompsis aggregata containing from 2 to 14 seeds. A paternity analysis based on ten polymorphic isozyme markers revealed multiple paternity in a minimum of 68% (based on simple paternity exclusion) and up to 100% (based on identification of the most likely father) of the multiseeded fruits. The estimated number of fathers increased with the number of seeds in a fruit, with an average of four sires, and up to nine sires, represented in a single fruit. To explore whether this level of multiple paternity could be explained solely by simultaneous deposition of a mixed pollen load, I constructed a computer simulation model based on previous measurements of movement patterns and pollen carryover by the hummingbird pollinators. Model predictions provided a good match to observed values for number of sires per fruit. Thus, the extensive pollen carryover in this species and consequent mixed pollen loads can explain the high levels of multiple paternity in natural populations.     

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.     

Primary pollinator exclusion has divergent consequences for pollen dispersal and mating in different populations of a bird‐pollinated tree

Pollination by nectarivorous birds is predicted to result in different patterns of pollen dispersal and plant mating compared to pollination by insects. We tested the prediction that paternal genetic diversity, outcrossing rate and realized pollen dispersal will be reduced when the primary pollinator group is excluded from bird‐pollinated plants. Pollinator exclusion experiments in conjunction with paternity analysis of progeny were applied to Eucalyptus caesia Benth. (Myrtaceae), a predominantly honeyeater‐pollinated tree that is visited by native insects and the introduced Apis mellifera (Apidae). Microsatellite genotyping at 14 loci of all adult E. caesia at two populations (n = 580 and 315), followed by paternity analysis of 705 progeny, revealed contrasting results between populations. Honeyeater exclusion did not significantly impact pollen dispersal or plant mating at Mount Caroline. In contrast, at the Chiddarcooping site, the exclusion of honeyeaters led to lower outcrossing rates, a threefold reduction in the average number of sires per fruit, a decrease in intermediate‐distance mating and an increase in near‐neighbour mating. The results from Chiddarcooping suggest that bird pollination may increase paternal genetic diversity, potentially leading to higher fitness of progeny and favouring the evolution of this strategy. However, further experimentation involving additional trees and study sites is required to test this hypothesis. Alternatively, insects may be effective pollinators in some populations of bird‐adapted plants, but ineffective in others.     

Evolutionary consequences of ecological factors: pollinator reliability predicts mating‐system traits of a perennial plant

The reproductive‐assurance hypothesis predicts that mating‐system traits will evolve towards increased autonomous self‐pollination in plant populations experiencing unreliable pollinator service. We tested this long‐standing hypothesis by assessing geographic covariation among pollinator reliability, outcrossing rates, heterozygosity and relevant floral traits across populations of Dalechampia scandens in Costa Rica. Mean outcrossing rates ranged from 0.16 to 0.49 across four populations, and covaried with the average rates of pollen arrival on stigmas, a measure of pollinator reliability. Across populations, genetically based differences in herkogamy (anther–stigma distance) were associated with variation in stigmatic pollen loads, outcrossing rates and heterozygosity. These observations are consistent with the hypothesis that, when pollinators are unreliable, floral traits promoting autonomous selfing evolve as a mechanism of reproductive assurance. Extensive covariation between floral traits and mating system among closely related populations further suggests that floral traits influencing mating systems track variation in adaptive optima generated by variation in pollinator reliability.     

Intrinsic and extrinsic factors acting on the reproductive process in alpine‐snowbed plants: roles of phenology,biological interaction,and breeding system

Pollinator activity and competition for pollinators lead to quantitative and qualitative pollen limitations on seed production and affect the reproductive success of plant species, depending on their breeding system (e.g., self‐compatibility and heterospecific compatibility) and genetic load (e.g., inbreeding depression and hybrid inviability). In alpine ecosystems, snowmelt regimes determine the distribution and phenology of plant communities. Plant species growing widely along a snowmelt gradient often grow with different species among local populations. Their pollinators also vary in their abundance, activity, and behavior during the season. These variations may modify plant–pollinator and plant–plant interactions. We integrated a series of our studies on the alpine dwarf shrub, Phyllodoce aleutica (Ericaceae), to elucidate the full set of intrinsic (species‐specific breeding system) and extrinsic factors (snow condition, pollinator activity, and interspecific competition) acting on their reproductive process. Seasonality of pollinator activity led to quantitative pollen limitation in the early‐blooming populations, whereas in the late‐blooming populations, high pollinator activity ensured pollination service, but interspecific competition for pollinators led to qualitative and quantitative pollen limitation in less competitive species. However, negative effects of illegitimate pollen receipt on seed‐set success might be reduced when cryptic incompatibility systems (i.e., outcross pollen grains took priority over self‐ and heterospecific pollen grains) could effectively prevent ovule and seed discounting. Our studies highlight the importance of species‐specific responses of plant reproduction to changing pollinator availability along environmental gradients to understand the general features of pollination networks in alpine ecosystems.     

Reproductive system and fitness of Vriesea friburgensis,a self‐sterile bromeliad species

Reproductive biology and plant fertility are directly related to many aspects of plant evolution and conservation biology. Vriesea friburgensis is an epiphytic and terrestrial bromeliad endemic to the Brazilian Atlantic rainforest. Hand‐pollination experiments were used to examine the reproductive system in a wild population of V. friburgensis. Plant fertility was assigned considering flower production, fruit and seed set, seed germination, and pollen viability. Self‐sterility observed from spontaneous selfing and manual self‐pollination treatments may be the consequence of late‐acting self‐incompatibility. Hand‐pollination results indicated no pollen limitation in the population studied. Floral biology features such as a few daily open flowers, nectar production, and sugar concentration corroborate hummingbirds as effective pollinators, although bees were also documented as pollinators. Components of fitness such as high flower, fruit, and seed production together with high seed and pollen viability indicate that this wild population is viable. From a conservation point of view, we highlight that this self‐sterile species depends on pollinator services to maintain its population fitness and viability through cross‐pollination. Currently, pollinators are not limited in this population of V. friburgensis. Conversely, the maintenance and continuous conservation of this community is essential for preserving this plant–pollinator mutualism.     

Patterns of multiple paternity in fruits of Mimulus ringens (Phrymaceae)

Multiply sired fruits provide unambiguous evidence that pollen from two or more donors was deposited on a stigma and successfully fertilized ovules. Such multiple paternity within fruits can have important consequences for both parental and offspring fitness, but little is known about the frequency of multiple paternity or the mechanisms causing it. In this study we quantify the extent of multiple paternity in replicate experimental arrays of Mimulus ringens (square-stem monkeyflower) and use observations of pollinator behavior to infer mechanisms generating multiply sired fruits. In each array, floral displays were trimmed to two, four, eight, or 16 flowers per plant to span the range of display sizes observed in nature. In our sample of 204 fruits, more than 95% had two or more outcross pollen donors. The number of sires per fruit averaged 4.63 ± 0.10 (mean ± 1 SE), including selfs, and did not vary significantly with floral display treatment. Patterns of bumble bee foraging, combined with limited pollen carryover, suggest that observed levels of multiple paternity cannot be fully explained by single probes that deposited mixed pollen loads. Multiple probes to flowers, each delivering pollen from 1-3 different sires, are more likely to have caused the observed patterns. These sequential visits may reduce the potential for pollen competition and female choice based on pollen tube growth rate.     

Pollination by sexual deception promotes outcrossing and mate diversity in self‐compatible clonal orchids

The majority of flowering plants rely on animals as pollen vectors. Thus, plant mating systems and pollen dispersal are strongly influenced by pollinator behaviour. In Australian sexually deceptive orchids pollinated by male thynnine wasps, outcrossing and extensive pollen flow is predicted due to floral deception, which minimizes multiple flower visitations within patches, and the movement of pollinators under mate‐search rather than foraging behaviours. This hypothesis was tested using microsatellite markers to reconstruct and infer paternity in two clonal, self‐compatible orchids. Offspring from naturally pollinated Chiloglottis valida and C. aff. jeanesii were acquired through symbiotic culture of seeds collected over three seasons. In both species, outcrossing was extensive (t_m = 0.924–1.00) despite clone sizes up to 11 m wide. The median pollen flow distance based on paternity for both taxa combined was 14.5 m (n = 18, range 0–69 m), being larger than typically found by paternity analyses in other herbaceous plants. Unexpectedly for orchids, some capsules were sired by more than one father, with an average of 1.35 pollen donors per fruit. This is the first genetic confirmation of polyandry in orchid capsules. Further, we report a possible link between multiple paternity and increased seed fitness. Together, these results demonstrate that deceptive pollination by mate‐searching wasps enhances offspring fitness by promoting both outcrossing and within‐fruit paternal diversity.     

Pollination biology of the hexaploid self‐compatible species Turnera velutina (Passifloraceae)

• The evolution of monomorphisms from heterostylous ancestors has been related to the presence of homostyly and the loss of self‐incompatibility, allowing the occurrence of selfing, which could be advantageous under pollinator limitation. However, flowers of some monomorphic species show herkogamy, attraction and rewarding traits that presumably favour cross‐pollination and/or a mixed mating system. This study evaluated the contributions of pollinators, breeding system and floral traits to the reproduction of Turnera velutina, a herkogamous monomorphic species.
• Floral visitors and frequency of visits were recorded, controlled hand cross‐pollinations were conducted under greenhouse and natural conditions, and individual variation in floral traits was characterised to determine their contribution to seed production.
• Apis mellifera was the most frequent floral visitor. Flowers presented approach herkogamy, high variation in nectar features, and a positive correlation of floral length with nectar volume and sugar concentration. Seed production did not differ between manual self‐ and cross‐pollinations, controls or open cross‐pollinations, but autonomous self‐pollination produced, on average, 82.74% fewer seeds than the other forms, irrespective of the level of herkogamy.
• Differences in seed production among autonomous self‐pollination and other treatments showed that T. velutina flowers depend on insect pollination for reproduction, and that approach herkogamy drastically reduced seed production in the absence of pollen vectors. The lack of differences in seed production from manual cross‐ and self‐pollinations suggests the possible presence of a mixed mating system in the studied population. Overall, this species was possibly derived from a distylous ancestor but appears fully capable of outcrossing despite being monomorphic.

     

Pollen transfer in fragmented plant populations: insight from the pollen loads of pollinators and stigmas in a mass‐flowering species

Pollinator and/or mate scarcity affects pollen transfer, with important ecological and evolutionary consequences for plant reproduction. However, the way in which the pollen loads transported by pollinators and deposited on stigmas are affected by pollination context has been little studied. We investigated the impacts of plant mate and visiting insect availabilities on pollen transport and receipt in a mass‐flowering and facultative autogamous shrub (Rhododendron ferrugineum). First, we recorded insect visits to R. ferrugineum in plant patches of diverse densities and sizes. Second, we analyzed the pollen loads transported by R. ferrugineum pollinators and deposited on stigmas of emasculated and intact flowers, in the same patches. Overall, pollinators (bumblebees) transported much larger pollen loads than the ones found on stigmas, and the pollen deposited on stigmas included a high proportion of conspecific pollen. However, comparing pollen loads of emasculated and intact flowers indicated that pollinators contributed only half the conspecific pollen present on the stigma. At low plant density, we found the highest visitation rate and the lowest proportion of conspecific pollen transported and deposited by pollinators. By contrast, at higher plant density and lower visitation rate, pollinators deposited larger proportion of conspecific pollen, although still far from sufficient to ensure that all the ovules were fertilized. Finally, self‐pollen completely buffered the detrimental effects on pollination of patch fragmentation and pollinator failure. Our results indicate that pollen loads from pollinators and emasculated flowers should be quantified for an accurate understanding of the relative impacts of pollinator and mate limitation on pollen transfer in facultative autogamous species.     

Floral contrivances and specialised pollination mechanism strongly influence mixed mating in Wrightia tomentosa (Apocynaceae)

• Reproductive success of a plant species is largely influenced by the outcome of mating pattern in a population. It is believed that a significantly larger proportion of animal‐pollinated plants have evolved a mixed‐mating strategy, the extent of which may vary among species. It is thus pertinent to investigate the key contributors to mating success, especially to identify the reproductive constraints in depauperate populations of threatened plant species.
• We examined the contribution of floral architecture, pollination mechanism and breeding system on the extent of outcrossing rate in a near‐threatened tree species, Wrightia tomentosa. The breeding system was ascertained from controlled pollination experiments. In order to determine outcrossing rate, 60 open‐pollinated progeny were analysed using an AFLP markers.
• Although the trees are self‐compatible, herkogamy and compartmentalisation of pollen and nectar in different chambers of the floral tube effectively prevent spontaneous autogamy. Pollination is achieved through specialised interaction with moths. Differential foraging behaviour of settling moths and hawkmoths leads to different proportions of geitonogamous and xenogamous pollen on the stigma. However, most open‐pollinated progeny were the result of xenogamy (outcrossing rate, tm = 0.68).
• The study shows that floral contrivances and pollination system have a strong influence on mating pattern. The differential foraging behaviour of the pollinators causes deposition of a mixture of self‐ and cross‐pollen to produce a mixed brood. Inbreeding depression and geitonogamy appear to play a significant role in sustaining mixed mating in this species.

     

High paternal diversity in the self-incompatible herb Arabidopsis halleri despite clonal reproduction and spatially restricted pollen dispersal

The number of sires fertilizing a given dam is a key parameter of the mating system in species with spatially restricted offspring dispersal, since genetic relatedness among maternal sibs determines the intensity of sib competition. In flowering plants, the extent of multiple paternity is determined by factors such as floral biology, properties of the pollen vector, selfing rate, spatial organization of the population, and genetic compatibility between neighbours. To assess the extent of multiple paternity and identify ecological factors involved, we performed a detailed study of mating patterns in a small population of a self-incompatible clonal herb, Arabidopsis halleri . We mapped and genotyped 364 individuals and 256 of their offspring at 12 microsatellite loci and jointly analysed the level of multiple paternity, pollen and seed dispersal, and spatial genetic structure. We found very low levels of correlated paternity among sibs ( P _full-sib = 3.8%) indicating high multiple paternity. Our estimate of the outcrossing rate was 98.7%, suggesting functional self-incompatibility. The pollen dispersal distribution was significantly restricted (mean effective pollen dispersal distance: 4.42 m) but long-distance successful pollination occurred and immigrating pollen was at most 10% of all pollination events. Patterns of genetic structure indicated little extent of clonal reproduction, and a low but significant spatial genetic structure typical for a self-incompatible species. Overall, in spite of restricted pollen dispersal, the multiple paternity in this self-incompatible species was very high, a result that we interpret as a consequence of high plant density and high pollinator service in this population.     

Pollinator activity can explain variation in outcrossing rates within individual trees

Abstract We tested the hypothesis that the previously recorded higher outcrossing rates and numbers of seeds per capsule from the upper, than from the lower, branches of trees of Eucalyptus globulus (Myrtaceae) is the result of greater pollinator activity in the upper parts of the canopy. Observations of bird pollinators on 23 trees, with flowers distributed evenly between the upper and lower halves of canopies, supported this hypothesis. Birds spent significantly more time foraging, and commenced foraging significantly more often, in the upper halves than in the lower halves of canopies. Flowers in the upper halves of E. globulus canopies would therefore be expected to receive more outcross‐pollen from bird pollinators because they are usually visited more often and would probably receive a greater ratio of outcross‐ to self‐pollen. We propose that such variation in pollinator activity and outcross‐pollen deposition results in different selective pressures on the mating system and pollination syndrome in different parts of the canopy. This may result in balanced selection for these traits, contributing to the maintenance of the mixed mating and generalized pollination systems of E. globulus.     

Nectar secretion dynamic links pollinator behavior to consequences for plant reproductive success in the ornithophilous mistletoe Psittacanthus robustus

The mistletoe Psittacanthus robustus was studied as a model to link flower phenology and nectar secretion strategy to pollinator behaviour and the reproductive consequences for the plant. The bright‐coloured flowers presented diurnal anthesis, opened asynchronously throughout the rainy season and produced copious dilute nectar as the main reward for pollinators. Most nectar was secreted just after flower opening, with little sugar replenishment after experimental removals. During the second day of anthesis in bagged flowers, the flowers quickly reabsorbed the offered nectar. Low values of nectar standing crop recorded in open flowers can be linked with high visitation rates by bird pollinators. Eight hummingbirds and two passerines were observed as potential pollinators. The most frequent flower visitors were the hummingbirds Eupetomena macroura and Colibri serrirostris, which actively defended flowering mistletoes. The spatial separation between anthers, stigma and nectar chamber promotes pollen deposition on flapping wings of hovering hummingbirds that usually probe many flowers per visit. Seed set did not differ between hand‐, self‐ and cross‐pollinated flowers, but these treatments set significantly more seeds than flowers naturally exposed to flower visitors. We suggest that the limitation observed in the reproductive success of this plant is not related to pollinator scarcity, but probably to the extreme frequency of visitation by territorial hummingbirds. We conclude that the costs and benefits of plant reproduction depend on the interaction strength between flowers and pollinators, and the assessment of nectar secretion dynamics, pollinator behaviour and plant breeding system allows clarification of the complexity of such associations.     

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.     

Florally rich habitats reduce insect pollination and the reproductive success of isolated plants

Landscape heterogeneity in floral communities has the potential to modify pollinator behavior. Pollinator foraging varies with the diversity, abundance, and spatial configuration of floral resources. However, the implications of this variation for pollen transfer and ultimately the reproductive success of insect pollinated plants remains unclear, especially for species which are rare or isolated in the landscape. We used a landscape‐scale experiment, coupled with microsatellite genotyping, to explore how the floral richness of habitats affected pollinator behavior and pollination effectiveness. Small arrays of the partially self‐compatible plant Californian poppy (Eschscholzia californica) were introduced across a landscape gradient to simulate rare, spatially isolated populations. The effects on pollinator activity, outcrossing, and plant reproduction were measured. In florally rich habitats, we found reduced pollen movement between plants, leading to fewer long‐distance pollination events, lower plant outcrossing, and a higher incidence of pollen limitation. This pattern indicates a potential reduction in per capita pollinator visitation, as suggested by the lower activity densities and richness of pollinators observed within florally rich habitats. In addition, seed production reduced by a factor of 1.8 in plants within florally rich habitats and progeny germination reduced by a factor of 1.2. We show this to be a consequence of self‐fertilization within the partially self‐compatible plant, E. californica. These findings indicate that locally rare plants are at a competitive disadvantage within florally rich habitats because neighboring plant species disrupt conspecific mating by co‐opting pollinators. Ultimately, this Allee effect may play an important role in determining the long‐term persistence of rarer plants in the landscape, both in terms of seed production and viability. Community context therefore requires consideration when designing and implementing conservation management for plants which are comparatively rare in the landscape.     

Influence of pollinator grooming on pollen‐mediated gene dispersal in Mimulus ringens (Phrymaceae)

Pollinator foraging patterns and the dynamics of pollen transport influence the quality and diversity of flowering plant mating opportunities. For species pollinated by grooming pollinators, such as bees, the amount of pollen carried between a donor flower and potential recipient flowers depends on how grooming influences pollen transfer. To investigate the relationship between grooming and pollen‐mediated gene dispersal, we studied bumblebee (Bombus fervidus) foraging behavior and resulting gene dispersal in linear arrays of Mimulus ringens. Each of the 14 plants in an array had a unique multilocus genotype, facilitating unambiguous assignment of paternity to 1050 progeny. Each plant was trimmed to a single flower so that pollinator movements could be linked directly to resulting gene dispersal patterns. Pollen‐mediated gene dispersal was very limited. More than 95% of the seeds sired by a donor flower were distributed over the first three recipient flowers in the visitation sequence. However, seeds were occasionally sired on flowers visited later in the pollinator's floral visitation sequence. Intensive grooming immediately following pollen removal from a donor flower significantly increased the decay rate of the donor flower's gene dispersal curve. These results suggest that the frequency and relative intensity of grooming can have significant effects on patterns of pollen‐mediated gene dispersal from individual pollen donors.     

Pollen limitation,fruit abortion,and autonomous selfing in three populations of the perennial herb Ruellia nudiflora

Multiple factors determine plant reproductive success and their influence may vary spatially. This study addresses several factors influencing female reproductive success in three populations of Ruellia nudiflora, specifically we: (i) determine if fruit set is pollen‐limited and if pollinator visitation rates are related to this condition; (ii) estimate fruit set via autonomous self‐pollination (AS) and relate it to the magnitude of herkogamy; and (iii) evaluate if fruit abortion is a post‐pollination mechanism that determines the magnitude of pollen limitation. At each site we marked 35 plants, grouped as: unmanipulated control (C) plants subjected to open pollination, plants manually cross‐pollinated (MP), and plants excluded from pollinators and only able to self‐pollinate autonomously (AS). Fruit set was greater for MP relative to C plants providing evidence for pollen limitation, while a tendency was observed for lower fruit abortion of MP relative to C plants suggesting that fruit set is influenced not only by pollen delivery per se, but also by subsequent abortion. In addition, although pollinator visits varied significantly among populations, the magnitude of pollen limitation did not, suggesting that pollinator activity was not relevant in determining pollen limitation. Finally, fruit set tended to decrease with the degree of herkogamy for AS plants, but this result was inconclusive. These findings have contributed to identify which factors influence reproductive success in populations of R. nudiflora, with potentially relevant implications for population genetic structure and mating system evolution of this species.     

Urbanization increases floral specialization of pollinators

Understanding how urbanization alters functional interactions among pollinators and plants is critically important given increasing anthropogenic land use and declines in pollinator populations. Pollinators often exhibit short‐term specialization and visit plants of the same species during one foraging trip. This facilitates plant receipt of conspecific pollen—pollen on a pollinator that is the same species as the plant on which the pollinator was foraging. Conspecific pollen receipt facilitates plant reproductive success and is thus important to plant and pollinator persistence. We investigated how urbanization affects short‐term specialization of insect pollinators by examining pollen loads on insects’ bodies and identifying the number and species of pollen grains on insects caught in urban habitat fragments and natural areas. We assessed possible drivers of differences between urban and natural areas, including frequency dependence in foraging, species richness and diversity of the plant and pollinator communities, floral abundance, and the presence of invasive plant species. Pollinators were more specialized in urban fragments than in natural areas, despite no differences in the species richness of plant communities across site types. These differences were likely driven by higher specialization of common pollinators, which were more abundant in urban sites. In addition, pollinators preferred to forage on invasive plants at urban sites and native plants at natural sites. Our findings reveal indirect effects of urbanization on pollinator fidelity to individual plant species and have implications for the maintenance of plant species diversity in small habitat fragments. Higher preference of pollinators for invasive plants at urban sites suggests that native species may receive fewer visits by pollinators. Therefore, native plant species diversity may decline in urban sites without continued augmentation of urban flora or removal of invasive species.