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.     

Pollination ecology of Isoglossa woodii,a long‐lived,synchronously monocarpic herb from coastal forests in South Africa

Synchronous monocarpy in long‐lived plants is often associated with pollination by wind, in part because infrequent mass flowering may satiate pollinators. Selfing in synchronous monocarps may provide reproductive assurance but conflict with the benefits of outcrossing, a key evolutionary driver of synchrony. We predicted that animal‐pollinated species with synchronous flowering would have unspecialised flowers and attract abundant generalised pollinators, but predictions for selfing and outcrossing frequencies were not obvious. We examined the pollination biology of Isoglossa woodii (Acanthaceae), an insect‐pollinated, monocarpic herb that flowers synchronously at 4–7‐year intervals. The most frequent visitor to I. woodii flowers was the African honeybee, Apis mellifera adansonii. Hand‐pollination failed to enhance seed production, indicating that the pollinators were not saturated. No seed was set in the absence of pollinators. Seed set was similar among selfed and outcrossed flowers, demonstrating a geitonogamous mixed‐mating strategy with no direct evidence of preferential outcrossing. Flowers contained four ovules, but most fruits only developed one seed, raising the possibility that preferential outcrossing occurs by post‐pollination processes. We argue that a number of the theoretical concerns about geitonogamous selfing as a form of reproductive assurance do not apply to a long‐lived synchronous monocarp such as I. woodii.     

The relative importance of birds and bees in the pollination of Metrosideros excelsa (Myrtaceae)

Exclusion experiments were used to assess the effect of different pollinator groups on outcrossing and seed production in Metrosideros excelsa. The main study site was Little Barrier Island, New Zealand where indigenous bird and native solitary bees are the main flower visitors. Our results showed that native birds were more important pollinators of M. excelsa than native bees. Seed production was much higher in open pollination than in two exclusion experiments where either birds were excluded and native bees only had access to flowers, or where all pollinators had been excluded. The number of fertile seeds per capsule was 45% higher after open pollination than in treatments with bee visitation only and 28% higher than in treatments where all flower visitors were excluded. Estimated outcrossing rates were significantly higher (t_m = 0.71) for open pollination in the upper canopy (>4 m above‐ground level) where bird visitation is presumed to be more frequent than for a treatment with native bee access only (t_m = 0.40). Our results also suggest that a large proportion of seeds (66%) arise from autonomous self‐pollination when all pollinators are excluded. In four trees of a modified mainland population with predominantly introduced birds and a mixture of introduced and native bees there was no decrease in seed production for the treatment allowing bee access only, indicating that – in contrast to native bees – honeybees may be more efficient pollinators of M. excelsa. Observation of the foraging behaviour of both groups of bees showed that native bees contact the stigma of flowers less frequently than honeybees. This is likely to be a consequence of their smaller body size relative to honeybees.     

Does restored plant diversity play a role in the reproductive functionality of Banksia populations?

Vegetation structure and plant species diversity of restoration sites are predicted to directly affect pollinator attraction, with potential impacts on gene flow, reproduction, genetic diversity of future generations, and ultimately restoration success. We compared Banksia attenuata R.Br. (Proteaceae) in a low species diversity restoration site and an adjacent natural remnant. We assessed fecundity genetic diversity in adult plants and their offspring, mating system parameters and pollen dispersal using paternity assignment. Results were compared to an earlier study of reproductive functionality within a high species diversity restoration site that was restored in a similar manner, enabling us to investigate any association between plant species diversity and fecundity. Seed set data indicated no significant differences between restored and adjacent natural sites; however, seed set data between restoration sites was significantly different (2.08 ± 0.39 and 6.89 ± 1.12, respectively). The mean number of fruits (follicles) per inflorescence was not significantly different between restoration sites. Genetic diversity of adult plants and their offspring were comparable in all sites. Higher allelic richness and genetic differentiation in one restored site reflected sourcing beyond local provenance. Low correlated paternity indicated high levels of multiple siring of seeds and paternity assignment demonstrated strong genetic connectivity between sites. Reproductive functionality, as measured by fecundity and genetic diversity in the offspring of B. attenuata, is resilient to low species diversity within a restored plant community. We consider our results in the context of establishing seed production areas (SPAs) that maximize the quantity and genetic quality of Banksia seeds for restoration.     

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.     

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.     

Sugar landscapes and pollinator‐mediated interactions in plant communities

Pollinator‐mediated interactions between plants can play an important role for the dynamics of plant communities. Pollination services depend on the abundance and the foraging behaviour of pollinators, which in turn respond to the availability and distribution of floral resources (notably nectar sugar). However, it is still insufficiently understood how the ‘sugar landscapes’ provided by flowering plant communities shape pollinator‐mediated interactions between multiple plant species and across different spatial scales. A better understanding of pollinator‐mediated interactions requires an integrative approach that quantifies different aspects of sugar landscapes and investigates their relative importance for pollinator behaviour and plant reproductive success. In this study, we quantified such sugar landscapes from individual‐based maps of Protea shrub communities in the Cape Floristic Region, South Africa. The 27 study sites of 4 ha each jointly comprise 127 993 individuals of 19 species. We analysed how rates of visitation by key bird pollinators and the seed set of plants respond to different aspects of sugar landscapes: the distribution of nectar sugar amounts, as well as their quality, taxonomic purity and phenology. We found that pollinator visitation rates strongly depended on phenological variation of site‐scale sugar amounts. The seed set of focal plants increased with nectar sugar amounts of conspecific neighbours and with site‐scale sugar amounts. Seed set increased particularly strongly if site‐scale sugar amounts were provided by plants that offer less sugar per inflorescence. These combined effects of the amount, quality, purity and phenological variation of nectar sugar show that nectar sugar is a common interaction currency that determines how multiple plant species interact via shared pollinators. The responses of pollinator‐mediated interactions to different aspects of this interaction currency alter conditions for species coexistence in Protea communities and may cause community‐level Allee effects that promote extinction cascades.     

Flower biology of the cactus Coryphantha elephantidens in the tropical dry forest of Central Mexico

Floral traits and sexual systems in angiosperms are strategies that enhance outcrossing within hermaphrodite flowers and among individuals in a population. Sexual systems with unisexual flowers have also evolved among angiosperms, resulting in sex specialization. Furthermore, the interaction of floral traits and floral visitors determines successful plant reproduction. Globose cacti are bee pollinated, and variation in the diversity of their pollinator assemblages is strongly associated with floral phenotype. Our objective was to describe the floral biology of the cactus Coryphantha elephantidens and to determine its relationship with pollinators. Floral traits were studied by direct observations in live and fixed flowers. The breeding system was determined using two estimators based on floral morphology: pollen grains to ovules per flower (P/O) ratio and outcrossing index. Pollination treatments were conducted to determine the mating system. Floral visitors were recorded using direct observation. Flowers of C. elephantidens are variable in color, protandric, herkogamous and nectarless. Estimators of the breeding system indicated xenogamy, which is consistent with the obligate outcrossing revealed by the pollination experiment. Thirty-seven percent of the plants have female flowers that do not produce pollen, making this population functionally gynodioecious. Both fruit and seed set were high compared to other globose cacti. Pollinators included eight species of native bees, a more diverse pollinator assemblage than other globose cacti. Given the high pressure on pollen due to functional gynodioecy, nectarless flowers, an outcrossing mating system, and the necessity of pollinators to set seeds, we concluded that native bees are highly efficient pollinators that play a crucial role in the sexual reproduction of C. elephantidens.     

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.     

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.     

Responses of nectar‐feeding birds to floral resources at multiple spatial scales

The responses of animal pollinators to the spatially heterogeneous distribution of floral resources are important for plant reproduction, especially in species‐rich plant communities. We explore how responses of pollinators to floral resources varied across multiple spatial scales and studied the responses of two nectarivorous bird species (Cape sugarbird Promerops cafer, orange‐breasted sunbird Anthobaphes violacea) to resource distributions provided by communities of co‐flowering Protea species (Proteaceae) in South African fynbos. We used highly resolved maps of about 125 000 Protea plants at 27 sites and estimated the seasonal dynamics of standing crop of nectar sugar for each plant to describe the spatiotemporal distribution of floral resources. We recorded avian population sizes and the rates of bird visits to > 1300 focal plants to assess the responses of nectarivorous birds to floral resources at different spatial scales. The population sizes of the two bird species responded positively to the amount of sugar resources at the site scale. Within sites, the effects of floral resources on pollinator visits to plants varied across scales and depended on the resources provided by individual plants. At large scales (radii > 25 m around focal plants), high sugar density decreased per‐plant visitation rates, i.e. plants competed for animal pollinators. At small scales (radii < 5 m around focal plants), we observed either competition or facilitation for pollinators between plants, depending on the sugar amount offered by individual focal plants. In plants with copious sugar, per‐plant visitation rates increased with increasing local sugar density, but visitation rates decreased in plants with little sugar. Our study underlines the importance of scale‐dependent responses of pollinators to floral resources and reveals that pollinators’ responses depend on the interplay between individual floral resources and local resource neighbourhood.     

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.

     

Bird and insect pollinators differ in specialization and potential pollination services along disturbance and resource gradients

Combined studies of the communities and interaction networks of bird and insect pollinators are rare, especially along environmental gradients. Here, we determined how disturbance by fire and variation in sugar resources shape pollinator communities and interactions between plants and their pollinating insects and birds. We recorded insect and bird visits to 21 Protea species across 21 study sites and for 2 years in Fynbos ecosystems in the Western Cape, South Africa. We recorded morphological traits of all pollinator species (41 insect and nine bird species). For each site, we obtained estimates of the time since the last fire (range: 2–25 calendar years) and the Protea nectar sugar amount per hectare (range: 74–62 000 g/ha). We tested how post-fire age and sugar amount influence the total interaction frequency, species richness and functional diversity of pollinator communities, as well as pollinator specialization (the effective number of plant partners) and potential pollination services (pollination service index) of insects and birds. We found little variation in the total interaction frequency, species richness and functional diversity of insect and bird pollinator communities, but insect species richness increased with post-fire age. Pollinator specialization and potential pollination services of insects and birds varied differently along the environmental gradients. Bird pollinators visited fewer Protea species at sites with high sugar amount, while there was no such trend for insects. Potential pollination services of insect pollinators to Protea species decreased with increasing post-fire age and resource amounts, whereas potential pollination services of birds remained constant along the environmental gradients. Despite little changes in pollinator communities, our analyses reveal that insect and bird pollinators differ in their specialization on Protea species and show distinct responses to disturbance and resource gradients. Our comparative study of bird and insect pollinators demonstrates that birds may be able to provide more stable pollination services than insects.     

Pollination biology of the endangered orchid Cypripedium japonicum in a fragmented forest of Japan

Pollination biology studies of the endangered orchid Cypripedium japonicum were conducted in its natural habitat using pollinator observation and hand‐pollination experiments. The observed fruit set was as follows: artificial outcross‐pollinated, 100%; artificial self‐pollinated, 100%; pollinator‐excluded, 0%; and emasculated flowers, 0%. These results show that this species, although self‐compatible, is neither autogamous nor agamospermous. The fruit set for open‐pollinated flowers was 14.9%, which suggests that the study population was subject to pollinator limitation. The nectarless flowers of C. japonicum were exclusively visited and pollinated by the queens of two bumblebee species (Bombus ardens and B. diversus diversus). It is probable that the nectarless flowers of C. japonicum attract pollinators through a generalized food deceptive system.     

Seed set variation in wild Clarkia populations: teasing apart the effects of seasonal resource depletion,pollen quality,and pollen quantity

In habitats where resource availability declines during the growing season, selection may favor early‐flowering individuals. Under such ephemerally favorable conditions, late‐blooming species (and individuals) may be particularly vulnerable to resource limitation of seed production. In California, a region prone to seasonal drought, members of the annual genus Clarkia are among the last to flower in the spring. We compared pollen limitation (PL) of seed set and outcrossing rates between early‐ and late‐flowering individuals in two mixed‐mating Clarkia taxa to detect whether flowering time is associated with changes in seed set due to resource depletion, PL, or increased selfing. In 2008–2010, we hand‐pollinated one flower on a total of 1855 individual plants either Early (near the onset of flowering) or Late (near the end of flowering) in the flowering season and compared seed set to adjacent, open‐pollinated flowers on the same stem. To assess the contribution of pollen quality to reproduction, we first (2008) used allozymes to estimate outcrossing rates of seeds produced by Early and Late open‐pollinated flowers. Second (2009), we conducted an anther‐removal experiment to estimate self‐pollen deposition. Seed set in Clarkia unguiculata was not pollen‐limited. Clarkia xantiana ssp. xantiana was pollen‐limited in 2008 and 2010, but not 2009. PL did not differ between Early and Late treatments. In both taxa, seed set of Early flowers was greater than Late flowers, but not due to PL in the latter. Reproduction was generally pollinator‐dependent. Most pollen deposition was xenogamous, and outcrossing rates were >0.7 – and similar between Early and Late periods. These results suggest that pollen receipt and pollen quality remain seasonally consistent. By contrast, the resources necessary to provision seeds decline, reducing the fitness benefits associated with resource allocation to ovules.     

Pollination syndromes in a specialised plant‐pollinator interaction: does floral morphology predict pollinators in Calceolaria?

Pollination syndromes are defined as suites of floral traits evolved in response to selection imposed by a particular group of pollinators (e.g., butterflies, hummingbirds, bats). Although numerous studies demonstrated their occurrence in plants pollinated by radically different pollinators, it is less known whether it is possible to identify them within species pollinated by one functional pollinator group. In such a framework, we expect floral traits to evolve also in response to pollinator subgroups (e.g., species, genera) within that unique functional group. On this, specialised pollination systems represent appropriate case studies to test such expectations. Calceolaria is a highly diversified plant genus pollinated by oil‐collecting bees in genera Centris and Chalepogenus. Variation in floral traits in Calceolaria has recently been suggested to reflect adaptations to pollinator types. However, to date no study has explicitly tested that observation. In this paper, we quantitatively test that hypothesis by evaluating the presence of pollination syndromes within the specialised pollination system formed by several Calceolaria and their insect pollinators. To do so, we use multivariate approaches and explore the structural matching between the morphology of 10 Calceolaria taxa and that of their principal pollinators. Our results identify morphological matching between floral traits related to access to the reward and insect traits involved in oil collection, confirming the presence of pollinator syndromes in Calceolaria. From a general perspective, our findings indicate that the pollination syndrome concept can be also extended to the intra‐pollinator group level.     

Unsuccessful introduced biocontrol agents can act as pollinators of invasive weeds: Bitou Bush (Chrysanthemoides monilifera ssp. rotundata) as an example

The extent of self‐compatibility and reliance on pollinators for seed set are critical determinants of reproductive success in invasive plant species. Seed herbivores are commonly used as biocontrol agents but may also act as flower visitors, potentially resulting in pollination. However, such contrasting or potentially counterproductive interaction effects are rarely considered or evaluated for biological control programs. We investigated the breeding system and pollinators of Bitou Bush (Chrysanthemoides monilifera ssp. rotundata), an invasive species in Australia that has been the subject of biocontrol programs since 1987. We found the species to be obligate outcrossing in all six populations tested. From 150 video hours, we found 21 species of potential pollinators, including Mesoclanis polana, the Bitou Seedfly, native to South Africa and released in Australia as a biocontrol agent in 1996. Mesoclanis polana transferred pollen to stigmas and was the most common pollinator (52% of pollinator visits), followed by the syrphid fly Simosyrphus grandicornis (9%) and introduced honeybee, Apis mellifera (6.5%). Fruit‐to‐flower ratios ranged from 0.12 to 0.45 and were highest in the population with the greatest proportion of Mesoclanis polana visits. In an experimental trial, outside the naturalized range, the native bee Homalictus sphecodoides and the native syrphid Melangyna viridiceps were the primary pollinators, and fruit‐to‐flower ratios were 0.35, indicating that Bitou Bush would have ready pollinators if its range expanded inland. Synthesis. Invasive Bitou Bush requires pollinators, and this is effected by a range of generalist pollinators in eastern Australia including the Bitou Seedfly, introduced as a biocontrol agent, and the major pollinator detected in this study. Fruit‐to‐flower ratios were highest when the Bitou Seedfly was in high abundance. This study underscores the importance of evaluating the pollination biology of invasive species in their native ranges and prior to the introduction of biocontrol agents.     

A Genetic Assessment of Ecological Restoration Success in Banksia attenuata

Rarely assessed in the success of ecological restoration projects is the maintenance of genetic variation in restored populations and, critically, their offspring. A founding population sourced from a limited genetic pool of nonlocal provenance seed can result in genetic bottlenecking and inbreeding, potentially reducing future population resilience and restoration success. We used microsatellite markers to assess the genetic variation of natural and restored populations, and their offspring, in Banksia attenuata R.Br. (Proteaceae), a keystone species of Banksia woodlands in south‐west Australia. Both natural and restored populations, and their offspring, displayed similarly high levels of heterozygosity (H_e range = 0.57–0.62) and allelic diversity (N_e range = 6.67–8.86) across 7 microsatellite loci. There was very weak population divergence (F_ST = 0.006) between the restored population and the adjacent natural population, indicating local provenance sourcing of seed. Genetic structuring within the natural population was weak, but detectable at 10 m and more strongly genetically structured than the restored population (Sp = 0.006 and 0.002, respectively). Complete outcrossing, low‐correlated paternity, and very low bi‐parental inbreeding were observed in both populations. Extensive pollen dispersal was observed within and among populations, with >50% of paternity assigned to sires beyond the local population. In a greenhouse experiment, differences in the overall performance of seedlings from natural and restored populations were negligible. Results indicate the successful genetic management of B. attenuata in this restoration project, from which general principles emphasizing the use of diverse local provenance seeds, genetic integration, and delivery of pollinator services are supported.     

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.     

Pollination and reproduction enhance the invasive potential of an early invader: the case of Lythrum salicaria (purple loosetrife) in South Africa

The potential of an alien plant to spread rapidly and colonize new habitat may be related to the mode of reproduction and the ability to attract pollinators. Most studies focus on widespread invasive plants, in which pollinators are rarely limiting. Here, we assess the ability of a recent invader in South Africa, the tristylous Lythrum salicaria to self-reproduce and whether this can explain the delay between introduction and spread. This study was conducted in one of the largest known populations (a total of 7 populations in South Africa) of L. salicaria in the Liesbeek river in the fynbos biome. We assessed the importance of pollinators and autonomous selfing in L. salicaria by comparing seed set between pollinator excluded and naturally pollinated flowers. Overall, 5 pollinators (4 native and 1 alien) were recorded with Cape honeybees and Africa Monarch butterflies the most prominent. Seed and fruit set were significantly higher in open pollinated flowers compared to pollinator excluded flowers. Also, seed and fruit set in pollinator excluded flowers were higher in long and medium morphs compared to short morphs. Germination was high for seeds from pollinator, but also from pollinator excluded treatments. This shows that L. salicaria in South Africa is self-compatible to some extent, but it is frequented by pollinators, significantly increasing seed production. Despite L. salicaria being tristylous, all 3 morphs are present in South Africa and with a huge seed production, this species has the potential to become a major invader of rivers and wetlands in South Africa.