Heterospecific Pollen Deposition Research Articles

Heterospecific pollen avoidance strategy prevails in the generalized plant-pollinator network on Yongxing Island.

Heterospecific pollen (HP) deposition varies widely among species in communities, which has been explicated by two adaptation strategies: HP avoidance and HP tolerance. Studies of the plant-pollinator network have uncovered that oceanic island communities are highly generalized and strongly connected. It remains unclear, however, which strategy prevails in such communities. We examined stigma pollen deposition on 29 plant species, and assessed patterns of HP load size and diversity in the Yongxing Island community. We assessed the effects of phenotypic specialization and species-level network structural properties of plant species on pollen deposition among species. The hypothesis of three accrual patterns of HP within species was tested by illustrating the relationship between conspecific pollen (CP) and HP receipt. Extensive variation occurred among species in HP receipt, while 75.9% of species received less than 10% HP and one species received more than 40% HP throughout the community. Flower size strongly drives the variation of HP receipt, while network structural properties had no effect on the pollen receipt. Nineteen species showed no relationship between the number of HP and CP loads, and they received smaller HP load sizes and lower HP proportions. Most plant species evolved HP avoidance strategy, and HP receipt was an occasional event for most plant species in the generalized community. HP and CP receipts are independent of each other in plant species with the HP avoidance mechanism. Our results highlight that plants in the generalized pollination system may preferentially select to minimize the HP load on stigmas.

Minimal impacts of invasive Scaevola taccada on Scaevola plumieri via pollinator competition in Puerto Rico

IntroductionScaevola taccada and Scaevola plumieri co-occur on shorelines of the Caribbean. Scaevola taccada is introduced in this habitat and directly competes with native dune vegetation, including S. plumieri, a species listed as locally endangered and threatened in Caribbean locations. This study addresses whether the invasive S. taccada also impacts the native S. plumieri indirectly by competing for pollinators and represents the first comparative study of insect visitation between these species.MethodsInsect visitation rates were measured at sites where species co-occur and where only the native occurs. Where species cooccur, insect visitors were captured, identified and analyzed for the pollen they carry. Pollen found on open-pollinated flowers was analyzed to assess pollen movement between the two species. We also compared floral nectar from each species by measuring volume, sugar content, and presence and proportions of amine group containing constituents (AGCCs).ResultsOur results demonstrate that both species share insect visitors providing the context for possible pollinator competition, yet significant differences in visitation frequency were not found. We found evidence of asymmetrical heterospecific pollen deposition in the native species, suggesting a possible reproductive impact. Insect visitation rates for the native were not significantly different between invaded and uninvaded sites, suggesting that the invasive S. taccada does not limit pollinator visits to S. plumieri. Comparisons of nectar rewards from the invasive and the native reveal similar volumes and sugar concentrations, but significant differences in some amine group containing constituents that may enhance pollinator attraction.ConclusionOur analysis finds no evidence for pollination competition and therefore S. taccada’s main impacts on S. plumieri are through competitive displacement and possibly through reproductive impacts as a consequence of heterospecific pollen deposition.

Heterospecific pollen deposition in understory plants of a forest in central Amazon

In co-flowering plant communities, pollinator movement can lead to the transfer of heterospecific pollen among species. This can affect the reproductive success of individuals, either by preventing conspecific pollen from reaching the stigma or by allelopathy, but little is known about this process in tropical areas. To analyze heterospecific pollen transfer and deposition, we investigated attributes related to pollinator attraction, such as corolla size, number of flowers per individual, floral arrangement, floral asymmetry, and distance from one flowering individual to another in a Central Amazon forest. The study was carried out in riparian plots of the Adolpho Ducke Forest Reserve during the rainy season. Stigmas and anthers were removed from all individual understory plants found in bloom in permanent plots. Slides were mounted with glycerinated gelatin, and all conspecific and heterospecific pollen grains were identified and counted. We then used a generalized linear mixed model framework to determine how the measured variables were related to the proportion of heterospecific pollen deposition (HPD) on stigmas. We found that the mean proportion of HPD in individuals was 4.7%, and 60% of individuals had no heterospecific pollen; from the 14 species studied, two had no heterospecific pollen. Species with wider flowers received less HPD, while species with more flowers per individual and with solitary flowers received more heterospecific pollen. Despite the significant variation in floral types in the Adolpho Ducke Reserve, there was a low percentage of heterospecific pollen deposition, probably due to specialized plant-pollinator interactions.

Within-bloom shift in abundance of a wild pollinator mediates pollen deposition rates to blueberry

Intra-seasonal variation in abiotic and biotic conditions can have profound consequences for pollinator community compositions and foraging movement, with flow-on effects upon pollination services. Yet, few studies have related such variations to pollination services in crop systems. In a cultivated highbush blueberry system with two primary pollinators — the managed European honey bee and a wild stingless bee species — we investigated how pollinator abundances, bee foraging behaviour, and con- and heterospecific stigmatic pollen loads changed over early, mid, and late blueberry blooming. Both con- and heterospecific stigmatic pollen loads declined following early bloom. This shift was associated with a decline in the abundance of stingless bees, whereas the abundance of honey bees only declined during late bloom. Simultaneously, honey bees were more likely to forage for blueberry pollen, and stigmatic pollen loads, relative to bee abundance, increased during late bloom. Although mixed pollen loads were common on pollinator bodies, especially on pollen foraging honey bees, heterospecific pollen deposition on blueberry stigmas was low. Given the similar effectiveness of honey bees and stingless bees as pollinators of blueberries, we contend that the observed seasonal variation in pollen deposition is likely caused by the decline in stingless bee abundances, as honey bees were not able to fully compensate for the loss of stingless bees during late bloom. Greater consideration of seasonal heterogeneity of pollinator abundance and behaviour, as part of pollination management plans, may aid in ensuring high pollination services throughout the entirety of crop bloom.

Limited reproductive interference despite high rates of heterospecific pollen transfer among co-occurring bat-pollinated Burmeistera.

Many tropical plants are bat-pollinated, but these mammals often carry copious, multispecific pollen loads making bat-pollinated plants susceptible to heterospecific pollen deposition and reproductive interference. We investigated pollen transfer between sympatric bat-pollinated Burmeistera species and their response to heterospecific pollen deposition from each other. We quantified conspecific and heterospecific pollen deposition for two populations of B. ceratocarpa, a recipient species in heterospecific pollen transfer interactions, that co-occur with different donor relatives (B. borjensis and B. glabrata). We then used a cross-pollination scheme using pollen mixtures to assess the species' responses to heterospecific pollen deposition in terms of fruit abortion and seed production. Burmeistera ceratocarpa received significantly more heterospecific pollen from its relatives at both sites than its own pollen was deposited on its relatives. However, heterospecific pollen deposition only affected seed production by B. borjensis and B. glabrata, but not by B. ceratocarpa, suggesting that early acting post-pollination barriers buffer the latter against reproductive interference. Crosses between sympatric and allopatric populations suggest that the study species are fully isolated in sympatry, while isolation between allopatric populations is strong but incomplete. We did not observe evidence of reproductive interference among our study species, because either heterospecific pollen deposition did not affect their seed production (B. ceratocarpa) or they receive heterospecific pollen only rarely (B. borjensis and B. glabrata). Frequent heterospecific pollen deposition might favor the evolution of barriers against foreign pollen (as in B. ceratocarpa) that alleviate the competitive costs of sharing low fidelity pollinators with co-occurring species.

Pollen as the link between floral phenotype and fitness

AbstractPollen plays a key role in plant reproductive biology. Despite the long history of research on pollen and pollination, recent advances in pollen‐tracking methods and statistical approaches to linking plant phenotype, pollination performance, and reproductive fitness yield a steady flow of exciting new insights. In this introduction to the Special Issue “Pollen as the Link Between Phenotype and Fitness,” we start by describing a general conceptual model linking functional classes of floral phenotypic traits to pollination‐related performance metrics and reproductive fitness. We use this model as a framework for synthesizing the relevant literature, highlighting the studies included in the Special Issue, and identifying gaps in our understanding and opportunities for further development of the field. The papers that follow in this Special Issue provide new insights into the relationships between pollen production, presentation, flower morphology, and pollination performance (e.g., pollen deposition onto stigmas), the role of pollinators in pollen transfer, and the consequences of heterospecific pollen deposition. Several of the studies demonstrate exciting experimental and analytical approaches that should pave the way for continued work addressing the intriguing role of pollen in linking plant phenotypes to reproductive fitness.

Do Silene species with exposed stigmas tolerate interference by heterospecific pollen?

Co-flowering species that have not evolved an avoidance mechanism may have tolerance to heterospecific pollen (HP) deposition as an adaptive strategy to minimize any deleterious effects of HP transfer, but empirical evidence for the tolerance hypothesis remains scarce. To estimate the potential effects of heterospecific pollen deposition (HPD) on female reproductive success, we counted conspecific (CP) and HP pollen grains deposited on stigmas and assessed subsequent seed set of both open- and hand-pollinated flowers in three co-flowering Silene species with exposed stigmas that usually received numerous HP grains on the elongated receptive area. The percentage of HP grains per flower (HP%) varied from 16.6% to 43.0% among three species. Silene chungtienensis had lower HP%, and the CP-HP relationship was neutral; S. gracilicaulis and S. yunnanensis had a relatively higher HP% with a positive CP-HP relationship. The effects of CP and HP number on natural seed set were positive for all three species, but HP% had stronger negative effects in S. chungtienensis and S. gracilicaulis. In hand-pollinated flowers of the three Silene species, seed set did not decrease with HP whether CP was in excess or insufficient, indicating no negative effects of HPD on seed production. Consistent with the tolerance hypothesis, our results indicated that species with higher HP interference are likely to be tolerant to an increase in HP%. These species with generalist-pollinated flowers and exposed large stigmas may benefit from an increase of conspecific pollen deposition, despite the associated increase in heterospecific pollen deposition.

Nototribic Pollination Reduces Heterospecific Pollen Deposition and Dampens Its Negative Impact on Seed Production in Calceolaria Species

Premise of research. Nototriby has evolved independently many times in different plant families, possibly for the advantage in the prevention of heterospecific pollen contact in pollinator-sharing environments. However, differences in pollen deposition and their consequences have received little attention, especially in highly specialized systems. Methodology. A systematic stigma collection was performed to estimate the proportion of heterospecific and conspecific pollen in two Calceolaria species with different pollination modes (i.e., nototribic and sternotribic) inhabiting a high mountain valley in Chile. Then, with the previous information, a manipulative pollen donor-receipt experiment was conducted to determine the impact of heterospecific pollen deposition in female reproductive success in both species (seed production). Pivotal results. The two Calceolaria species were prone to receive pollen from some plants of the surrounding community, mainly Adesmia emarginata. However, in the nototribic species (C. filicaulis), only 10% of the total pollen deposition belonged to heterospecific plant species, while in the sternotribic species (C. cavanillesii), more than 60% was heterospecific. Concomitant with this, the hand cross-pollination experiment showed that seed production of C. cavanillesii was more affected by A. emarginata pollen deposition than C. filicaulis pollen. Conclusions. Nototribic morphology can be an effective prepollination barrier to the deposition of heterospecific pollen. However, when this barrier is overcome, its impact may be affected by different factors, such as the foraging behavior of their main pollinator species and a rejection or tolerance mechanism to foreign pollen receipt developed by plant species. Future studies should consider assays to determine the mechanisms behind these findings and their generality to other species of the genus Calceolaria.

Conspecific pollen advantage mediated by the extragynoecial compitum and its potential to resist interspecific reproductive interference between two Sagittaria species.

The extragynoecial compitum formed by the incomplete fusion of carpel margins, while allowing intercarpellary growth of pollen tubes in apocarpous angiosperms, may also increase the risk of reproductive interference caused by heterospecific pollen (HP) deposition. In Sagittaria, congeneric HP tubes grow via different paths and enter the ovules later than conspecific pollen (CP) tubes. However, it is unclear how the growth advantage of the CP tube helps ensure reproductive success when HP is deposited on the stigmas. We performed molecular characterization of interspecies-pollinated seeds to examine the consequences of interspecific pollen deposition between Sagittaria pygmaea and S. trifolia. We also conducted CP–HP (1:1) mixed pollination and delayed CP pollination treatments to explore the seed-siring abilities of CP and HP. Our results showed that although HP could trigger the development of fruits, the interspecies-pollinated seeds contained partially developed embryos and could not germinate. More than 70% of the embryos in these seeds were molecularly identified as hybrids of both species, suggesting that HP tubes could enter the ovules and fertilize the egg cells. Moreover, CP could sire more offspring (≥70%) after the CP–HP (1:1) mixed pollination treatment, even when HP reached the stigma 0.5–1 h earlier than CP (≥50%). Following adequate CP vs. HP (1:1) pollination on carpels on two sides of the apocarpous gynoecium, both species produced > 70% conspecific seeds, indicating that the CP tubes could occupy ovules that should be occupied by HP via the extragynoecial compitum. Our results reveal that in Sagittaria, pollen deposition from co-existing congeneric heterospecies leads to interspecific seed discounting. However, the CP advantage mediated by the extragynoecial compitum is an effective strategy to mitigate the effects of interspecific pollen deposition. This study improves our understanding of how apocarpous angiosperms with an extragynoecial compitum can maintain species stability and mitigate the negative reproductive interference effect from sympatrically distributed related species.

Hummingbird contribution to plant reproduction in the rupestrian grasslands is not defined by pollination syndrome.

Floral traits mediate the roles of distinct animals as effective pollinators along a generalization/specialization continuum. Many plant species are visited by different pollinator functional groups and the specific contribution of each group is expected to reflect the set of floral characteristics defined by pollination syndromes. Although considered a highly specialized nectarivorous group, hummingbirds frequently visit flowers lacking apparent specialization to bird pollination. How they contribute to the reproduction of these plants, however, has not been evaluated through field experiments considering multiple non-related plant species simultaneously. Here, we investigated hummingbirds' contributions to the pollination of ten plant species comprising a gradient of adaptation to bird pollination in the Brazilian rupestrian grasslands. We excluded hummingbirds from flowers and evaluated their relative contribution in comparison to insects (mainly bees) on conspecific/heterospecific pollen deposition and fruit set. Floral traits that are typically associated with bird pollination were associated with increased pollen deposition, but not with fruit set in the presence of hummingbirds. With hummingbirds, conspecific and heterospecific pollen deposition increased in most species, while fruit set increased in four plant species with varying degrees of fit to ornithophily. Our results show that assessing the relative contribution of specific pollinator groups may depend on when this contribution is measured, i.e. pollen deposition or fruit set. Considering fruit set, our results indicate that hummingbirds contributed to plant reproduction independently of the fit to bird pollination syndrome. This emphasizes their importance as under-appreciated generalized pollinators in some communities.

Heterospecific pollen deposition is positively associated with reproductive success in a diverse hummingbird‐pollinated plant community

Heterospecific pollen deposition (HPD) is ubiquitous across plant communities, especially for generalized species which use a diversity of pollinators, and may have negative effects on plant reproduction. However, it is unclear whether temporal changes in the co‐flowering community result in changes in HPD patterns. Moreover, community‐level studies are required to understand which factors influence HPD and how the reproduction of different species is affected.We investigated the temporal variation of HPD, its relationship with level of specialization on pollinators and floral phenotypic specialization, and its association with reproductive success (pollen limitation and fruit set) in 31 hummingbird‐pollinated plant species in a tropical Campo Rupestre.We found seasonality in HPD, with species flowering in the dry season having greater diversity of heterospecific pollen on stigmas and a higher frequency of stigmas containing heterospecific pollen, compared to the rainy season. Stigmas of ecologically generalized species had more heterospecific pollen, while the relationship for ecologically specialized species depended on floral phenotype. Surprisingly, and in contrast to theory, we found a positive relationship between HPD and reproductive success.Our results indicate benefits of generalization and facilitation, in which sharing pollinators brings greater reproductive success via increased conspecific pollen deposition, even if it incurs more HPD. We demonstrated how assessing HPD at a community‐level can contribute to understanding the ecological causes and functional consequences of pollinator sharing.

Spatial variation in bidirectional pollinator-mediated interactions between two co-flowering species in serpentine plant communities.

Pollinator-mediated competition and facilitation are two important mechanisms mediating co-flowering community assembly. Experimental studies, however, have mostly focused on evaluating outcomes for a single interacting partner at a single location. Studies that evaluate spatial variation in the bidirectional effects between co-flowering species are necessary if we aim to advance our understanding of the processes that mediate species coexistence in diverse co-flowering communities. Here, we examine geographic variation (i.e. at landscape level) in bidirectional pollinator-mediated effects between co-flowering Mimulus guttatus and Delphinium uliginosum. We evaluated effects on pollen transfer dynamics (conspecific and heterospecific pollen deposition) and plant reproductive success. We found evidence of asymmetrical effects (one species is disrupted and the other one is facilitated) but the effects were highly dependent on geographical location. Furthermore, effects on pollen transfer dynamics did not always translate to effects on overall plant reproductive success (i.e. pollen tube growth) highlighting the importance of evaluating effects at multiple stages of the pollination process. Overall, our results provide evidence of a spatial mosaic of pollinator-mediated interactions between co-flowering species and suggest that community assembly processes could result from competition and facilitation acting simultaneously. Our study highlights the importance of experimental studies that evaluate the prevalence of competitive and facilitative interactions in the field, and that expand across a wide geographical context, in order to more fully understand the mechanisms that shape plant communities in nature.

Comparisons among populations and individuals to evaluate pollen-pistil interaction as a mechanism of reproductive interference in Taraxacum.

Reproductive interference (RI), an interspecific mating interaction that reduces the fitness of at least one of the species involved, can lead to exclusive distributions in closely related species. A hypothesis previously proposed is that RI in plants may occur by ovule usurpation, in which pistils lack interspecific incompatibility and mistakenly accept heterospecific pollen, thereby losing an opportunity for conspecific pollen fertilization. However, few comparative studies have evaluated the consistency of the inferred mechanism within and among individuals and populations. We conducted hand-pollination experiments in six populations of three native Taraxacum species that suffered from different levels of RI from an alien congener, T. officinale, and compared pollen-pistil interactions among populations. We also investigated the interactions for eight individual T. japonicum plants whose response to heterospecific pollen deposition had been previously measured. Our results revealed that pollen tubes often penetrated native ovaries following heterospecific pollination in populations suffering from strong RI, whereas they seldom did in populations suffering from marginal RI. However, the relative frequency of the pollen tube penetration was not significantly related to the strength of alien RI. Not all pistils on an individual plant showed the same pollen receptivity following heterospecific pollination; rather, some accepted and some refused the pollen tubes. The relationship between pollen tube penetration following heterospecific pollination and the strength of the alien RI was also not significant among individuals. Our present results generally support the ovule usurpation hypothesis, but suggest that other factors, such as competition for pollinator services, variation in the effects of heterospecific pollen donors, and condition of the native inflorescences, might also affect the observed RI strength.

Coflowering invasive plants and a congener have neutral effects on fitness components of a rare endemic plant.

Network analyses rarely include fitness components, such as germination, to tie invasive plants to population‐level effects on the natives. We address this limitation in a previously studied network of flower visitors around a suite of native and invasive plants that includes an endemic plant at Badlands National Park, South Dakota, USA. Eriogonum visheri coflowers with two abundant invasive plants, Salsola tragus and Melilotus officinalis, as well as a common congener, E. pauciflorum. Network analyses had suggested strong linkages between E. visheri and S. tragus and E. pauciflorum, with a weaker link to M. officinalis. We measured visitation, pollen deposited on stigmas, achene weight and germination over three field seasons (two for germination) in four populations (two in the final season) of E. visheri and applied in situ pollen treatments to E. visheri, adding pollen from other flowers on the same plant; flowers on other E. visheri plants; S. tragus, M. officinalis, or E. pauciflorum; open pollination; or excluding pollinators. Insect visitation to E. visheri was not affected by floral abundance of any of the focal species. Most visitors were halictid bees; one of these (Lasioglossum packeri) was the only identified species to visit E. visheri all three years. Ninety‐seven percent of pollen on collected E. visheri stigmas was conspecific, but 22% of flowers had >1 grain of E. pauciflorum pollen on stigmas and 7% had >1 grain of S. tragus pollen; <1% of flowers had M. officinalis pollen on stigmas. None of the pollen treatments produced significant differences in weight or germination of E. visheri achenes. We conclude that, in contrast to the results of the network analysis, neither of the invasive species poses a threat, via heterospecific pollen deposition, to pollination of the endemic E. visheri, and that its congener provides alternative pollen resources to its pollinators.

The influence of floral resources and microclimate on pollinator visitation in an agro-ecosystem

As agriculture expands to meet the needs of a growing global population, natural ecosystems are threatened by deforestation and habitat fragmentation. Tropical agroforestry systems offer a sustainable alternative to traditional agriculture by providing food for production while also supporting biodiversity and ecosystem services. Previous studies have shown that these systems may even improve crop pollination, but the mechanisms of how these improvements occur are still poorly understood. Using coffee as a focal crop, we explored how microclimatic conditions affected nectar traits (sugar and caffeine concentration) important for pollinator visitation. We also studied how microclimate, floral traits, floral availability at the coffee plant level, availability of floral resources provided by other plant species in the agroecosystem (“neighborhood floral availability”), and the presence of other bees affected the amount of time bees spent foraging on coffee flowers and the proportion of coffee pollen carried on their bodies. We explored these factors using the two dominant coffee species farmed on Puerto Rico, Coffea canephora and C. arabica, under sun and shade management. We found that high nectar sugar concentration and temperature were important predictors of short floral visits (<15s), while increased numbers of bees and open coffee flowers were important predictors of longer floral visits (16–180 seconds). High nectar caffeine concentration was an important predictor of longer visits on C. arabica flowers while the opposite was observed for C. canephora flowers. For both species, high coffee floral availability was the main predicting factor for the proportion of coffee pollen on the bees’ bodies. Surprisingly, neither neighborhood floral availability nor the type of coffee plantation (agroforest/shade or sun) were important predictors of bee visitation. These results suggest non-coffee flowering plants in coffee plantations were neither competitors nor facilitators of coffee plants for pollinators. Additionally, most of the bees surveyed were carrying ≥80 % pollen from one species (C. arabica or C. canephora), likely resulting in little heterospecific pollen deposition between Coffea and non-Coffea flowers. Shade trees in coffee plantations do not detract from pollinator visitation to coffee flowers, suggesting that the provision of multiple ecological and wildlife conservation benefits by shade trees is not in conflict with a grower’s ability to maximize the benefits of insect pollination on fruit production.

Foraging behavior and pollination efficiency of generalist insects in an understory dioecious shrub Helwingia japonica

It has been hypothesized that pollination success in animal-pollinated dioecious plants relies on opportunistic pollinators with no discrimination against female flowers. However, empirical studies of pollinator foraging behavior and pollination effectiveness in dioecious species are few. To investigate potential pollinators in Helwingia japonica, a dioecious shrub with small, inconspicuous flowers, we compared floral visitors and visit frequency to female and male plants for three flowering seasons in two field populations in subtropical forests in southwest China. Pollen placement on the insect bodies of four groups (solitary bees and other bees, fungus gnats, and other flies) was compared, and insect foraging behavior was observed. Pollen removal and conspecific and heterospecific pollen deposition per visit were measured to compare pollination effectiveness among the four groups. Floral visitors usually did not discriminate between male and female flowers and did not gather pollen into collections. Our measurements of pollen transfer efficiency showed that solitary bees were the most effective pollinators with the highest conspecific pollen deposition. These insects seemed to be opportunistic visitors because pollen grains of H. japonica were distributed evenly over different regions of the visitor's body, and heterospecific pollen accounted for over 50% of total pollen loads on stigmas in the two populations. Our investigations indicated that potential pollinators were generalists and did not discriminate against female flowers, as predicted for dioecious species pollinated by insects. A perspective of pollen removal by floral visitors offers insights into the evolution of plant sexual systems.

Impact of Invasive Bees on Plant-Pollinator Interactions and Reproductive Success of Plant Species in Mixed Nothofagus antarctica Forests.

Invasive social bees can alter plant-pollinator interactions with detrimental effects on both partners. However, most studies have focused on one invasive bee species, while the interactions among two or more species remain poorly understood. Also, many study sites had a history of invasive bees, being hard to find sites with historical low abundances. In Patagonia, Bombus ruderatus (F.) invasion begun in 1993 and B. terrestris (L.) in 2006. Though honey bees (Apis mellifera L.) introduction started in 1859, their density is still low in some parts. By experimentally increasing honey bee densities, we evaluated the effect of honey bees and bumblebees floral visitation on native pollinator floral visitation, pollen deposition, and reproductive success of three plant species in mixed Nothofagus antarctica forests of northern Patagonia: Oxalis valdiviensis, Mutisia spinosa and Cirsium vulgare. Our results show that exotic bees became the main floral visitors. No negative association was found between invasive bee and native pollinator visitation rates, but there was evidence of potential competition between honey bees and bumblebees. Floral neighborhood diversity played an important role in pollinator behavior. Conspecific pollen deposition was high for all species, while deposition of heterospecific pollen was very high in M. spinosa and C. vulgare. Not as expected, honey bees visitation rate had a negative effect on heterospecific pollen deposition in C. vulgare. For O. valdiviensis, exotic visitation rates increased conspecific pollen deposition, which was positively related to reproductive success. Although exotic bees became main floral visitors, their contribution to reproductive success was only clear for one species.

Flower constancy of the Cape honey bee pollinator of two co-flowering Erica species from the Cape Floristic Region (South Africa)

Erica is the largest genus of the Cape Floristic Region (CFR) of South Africa, and multiple species often coexist. Species that co-flower and share a pollinator may suffer from reproductive interference through heterospecific pollen deposition, especially when such species are closely related. However, little is known about the mechanisms that enable coexistence of co-flowering Erica species that share pollinators. Here we focus on a site where two co-flowering Erica species which are both predominantly pollinated by Cape honey bees grow intermingled, to ask whether this is associated with flower constancy and/or differences in pollen deposition sites. Despite minor differences in sites of pollen deposition on honey bees, most pollen of both Erica species was found on the head and ventral thorax regions. Subsequent visits of honey bees were almost always to the same Erica species from which they departed, and were significantly more constant than expected from random foraging in two consecutive years. Based on differences in pollen types between the two species, we established that pollen loads carried by honey bees were between 67% and 99% species-specific, whereas pollen loads on stigmas were on average over 93% pure. We conclude that despite the superficial similarities of small-flowered Erica species, honeybees can use species-specific cues for constant foraging, which is associated with almost complete absence of heterospecific pollen transfer. Future analyses of pollen carryover rates could help to quantify the benefit of pollinator behaviour for plant fecundity, and inclusion of more study sites sampled throughout the Erica flowering season could shed further light on the importance of flower constancy for facilitating coexistence of Erica species in the CFR.

Effect of heterospecific pollen deposition on pollen tube growth depends on the phylogenetic relatedness between donor and recipient.

Co-flowering plant species may interact via pollinators leading to heterospecific pollen transfer with consequences for plant reproduction. What determines the severity of heterospecific pollen effect on conspecific pollen performance is unclear, but it may depend on the phylogenetic relatedness of the interactors (pollen donors and recipient). The heterospecific pollen effect might also depend on the extent to which plants are exposed to heterospecific pollen over ecological or evolutionary timescales. For instance, generalist-pollinated plant species might tolerate heterospecific pollen more than specialists. Here, we tested whether heterospecific pollen effects are stronger between closely related species than phylogenetically distant ones in a tropical highland community. Then, based on these results, we determined whether responses to heterospecific pollen were stronger in generalized vs. specialized plant species. We applied heterospecific pollen from close (congeneric) or distant (different families) donors alone or with conspecific pollen on stigmas of three recipient species (one generalist, Sisyrinchium wettsteinii; and two specialists, Fuchsia campos-portoi and Fuchsia regia) and scored pollen tube performance in styles. In all species, pollen from closely related donors grew pollen tubes to the base of the style indicating a high potential to interfere with seed set. Conversely, distantly related heterospecific pollen had no effect on either specialist Fuchsia species, whereas enhanced performance of conspecific pollen was observed in generalist S. wettsteinii. The strong effect of phylogenetic relatedness of donor and recipient might have obscured the role of pollination specialization, at least for the three species examined here. Therefore, phylogenetic relatedness mediated the effect of heterospecific pollen on post-pollination success, with possible consequences for reproductive trait evolution and community assembly for further studies to explore.

A new native plant in the neighborhood: effects on plant-pollinator networks, pollination, and plant reproductive success.

Ecological communities are dynamic entities subjected to extinction/colonization events. Because species are connected through complex interaction networks, the arrival of a new species is likely to affect various species across the community, as observed in plant biological invasions. However, plant invasions usually represent extreme scenarios in which the community is strongly dominated by the alien species, confounding the effects of a change in species composition with a massive increase in floral resource availability. Our study addresses changes in plant community composition involving native species, a common phenomenon under the current climate change scenario in which plants are modifying their distribution ranges. We experimentally manipulated patches of a natural scrubland community by introducing a native plant (henceforth colonizing plant). To avoid introducing a disproportionate amount of floral resources we adjusted the number of flowers of the colonizing plant to the amount of floral resources locally available in each patch. We had two objectives: (1) to analyse the effects of the arrival of a new plant on the pollinator community, the rearrangement of plant-pollinator interactions and the structure of the plant-pollinator network; (2) to evaluate potential consequences for pollination and the reproductive success of resident plant species. The colonizing plant acted as a magnet species, attracting bumble bees and facilitating interactions to other plants through spill-over. The introduction of the colonizing plant also affected the structure of plant-pollinator networks (colonized networks were more generalized and more nested than control networks) and modified the arrangement of plant and pollinator species into modules. Ultimately, these changes resulted in higher heterospecific (but not conspecific) pollen deposition and had contrasting effects on the reproductive success of two resident plant species (higher fruit set and lower seed set, respectively). Our study shows that relationships between plants and pollinators are rapidly rearranged in response to novel situations (even when the new plant is not overly dominant), with important functional consequences on pollination and plant reproductive success. Our study establishes a link between network structure and pollination and plant reproductive success, which may be mediated by differences among pollinator species in foraging behavior.