The ecological dynamics of co-flowering communities are largely mediated by pollinators. However, current understanding of pollinator-mediated interactions primarily relies on how co-flowering plants influence attraction of shared pollinators, and much less is known about plant–plant interactions that occur via heterospecific pollen (HP) transfer. Invaded communities in particular can be highly affected by the transfer of alien pollen, but the strength, drivers and fitness consequences of these interactions at a community scale are not well understood. Here we analyse HP transfer networks in nine coastal communities in the Yucatan Mexico that vary in the relative abundance of invasive flowers to evaluate how HP donation and receipt varies between native and alien plants. We further evaluate whether HP donation and receipt are mediated by floral traits (e.g. display, flower size) or pollinator visitation rate. Finally, we evaluated whether post-pollination success (proportion of pollen tubes produced) was affected by alien HP receipt and whether the effect varied between native and alien recipients. HP transfer networks exhibit relatively high connectance (c. 15%), suggesting high HP transfer within the studied communities. Significant network nestedness further suggests the existence of species that predominantly act as HP donors or recipients in the community. Species-level analyses showed that natives receive 80% more HP compared to alien species, and that alien plants donate 40% more HP than natives. HP receipt and donation were mediated by different floral traits and such effects were independent of plant origin (native or alien). The proportion of alien HP received significantly affected conspecific pollen tube success in natives, but not that of alien species. Synthesis. Our results suggest that HP transfer in invaded communities is widespread, and that native and alien species play different roles within HP transfer networks, which are mediated by a different suite of floral traits. Alien species, in particular, play a central role as HP donors and are more tolerant to HP receipt than natives—a finding that points to two overlooked mechanisms facilitating alien plant invasion and success within native co-flowering communities.

The number of co-flowering species, floral density and floral trait diversity can be major determinants of pollinator-mediated plant–plant interactions in a community. However, evaluation of how each one of these co-flowering components affects the pollination success of a single focal plant species, and how these effects vary at different spatial scales, is lacking. Here, we evaluated the effects of functional diversity (flower morphology and colour), taxonomic diversity (reflecting potential sampling effects) and flower density (conspecific and heterospecific), on the pollinator environment (i.e. visitation rate and pollinator diversity) and pollination success (i.e. pollen load size and number of pollen tubes per style) of Cakile edentula (Brassicaceae). We applied structural equation models (SEMs) at the floral-neighbourhood (plot level) and community-wide scales to uncover the factors that mediate co-flowering community effects on C. edentula pollination success. We found that co-flowering community effects at the community level are more important than fine-scale floral-neighbourhood differences in mediating plant pollination success in our study species. Increasing plant functional diversity decreased pollinator visitation rate but increased the diversity of pollinator functional groups visiting C. edentula flowers. Taxonomic diversity negatively affected pollinator diversity suggesting that other unmeasured floral traits may be relevant or that single-species effects (sampling effects) may be important. Overall, our results suggest that functional floral trait diversity in a community may be the most important factor influencing pollination success of species in a community. We also found evidence for intra- and interspecific pollinator competition mediated by flower density, but none of these effects seemed to have a significant impact on pollination success. This study is an important step towards understanding the complexity of co-flowering community effects on the pollination success of individual plant species at multiple spatial scales. This study further reveals the potential importance of plant functional diversity in a community in helping predict competitive and facilitative interactions in co-flowering communities. Synthesis. Floral density and taxonomic and functional co-flowering diversity are important drivers of pollination success in flowering plants. The effects of the co-flowering diversity on the pollination success of plant species can largely depend on the spatial scale being studied. Only evaluating the outcomes of pollinator-mediated plant–plant interactions at multiple stages of the pollination process can lead to a complete understanding of their ecological consequences in nature.

Premise of The Study: Studies on the effects of heterospecific pollen (HP) transfer have been focused mainly on insect-pollinated species, despite evidence of insect visitation to wind-pollinated species and transfer of their pollen onto stigmas of insect-pollinated plants. Thus, the potential consequences of HP transfer from wind-pollinated species remain largely unknown. Furthermore, accumulation of pesticide residues in pollen of wind-pollinated crops has been documented, but its potential effects on wild plant species via HP transfer have not been tested. Methods: We evaluated the effect of wind-dispersed Zea mays pollen on pollen tube growth of the insect-pollinated Mimulus nudatus via hand pollinations. We further evaluated whether pesticide-contaminated Z. mays pollen has larger effects on M. nudatus pollen success than non-contaminated Z. mays pollen. Key Results: We found a significant negative effect of Z. mays pollen on M. nudatus pollen tube growth even when deposited in small amounts. However, we did not observe any difference in the magnitude of this effect between pesticide-laden Z. mays pollen and non-contaminated Z. mays pollen. Conclusions: Our results suggest that wind-pollinated species can have negative effects as HP donors on insect-pollinated recipients. Thus, their role in shaping co-flowering interactions for wind- and insect-pollinated species deserves more attention. Although we did not find evidence that pesticide contamination increased HP effects, we cannot fully rule out the existence of such an effect, because pollen load and thus the pesticide dose applied to stigmas was low. This result should be confirmed using other HP donors and across a range of HP loads, pesticide types, and concentrations.

The ecological dynamics of co-flowering communities are largely mediated by pollinators. However, current understanding of pollinator-mediated interactions primarily relies on how co-flowering plants influence attraction of shared pollinators, and much less is known about plant–plant interactions that occur via heterospecific pollen (HP) transfer. Invaded communities in particular can be highly affected by the transfer of alien pollen, but the strength, drivers and fitness consequences of these interactions at a community scale are not well understood. Here we analyse HP transfer networks in nine coastal communities in the Yucatan Mexico that vary in the relative abundance of invasive flowers to evaluate how HP donation and receipt varies between native and alien plants. We further evaluate whether HP donation and receipt are mediated by floral traits (e.g. display, flower size) or pollinator visitation rate. Finally, we evaluated whether post-pollination success (proportion of pollen tubes produced) was affected by alien HP receipt and whether the effect varied between native and alien recipients. HP transfer networks exhibit relatively high connectance (c. 15%), suggesting high HP transfer within the studied communities. Significant network nestedness further suggests the existence of species that predominantly act as HP donors or recipients in the community. Species-level analyses showed that natives receive 80% more HP compared to alien species, and that alien plants donate 40% more HP than natives. HP receipt and donation were mediated by different floral traits and such effects were independent of plant origin (native or alien). The proportion of alien HP received significantly affected conspecific pollen tube success in natives, but not that of alien species. Synthesis. Our results suggest that HP transfer in invaded communities is widespread, and that native and alien species play different roles within HP transfer networks, which are mediated by a different suite of floral traits. Alien species, in particular, play a central role as HP donors and are more tolerant to HP receipt than natives—a finding that points to two overlooked mechanisms facilitating alien plant invasion and success within native co-flowering communities.

Premise of research. The duration and intensity of flowering overlap among plants are the first determiners of the potential for pollinator-mediated plant-plant interactions. Yet, our ability to describe community-wide patterns of coflowering, and thus understand its impact on the structure of plant-pollinator communities, is limited. Methodology. We present a conceptual framework for how network theory can reveal structural properties that are ecologically relevant in diverse coflowering communities. Coflowering modules, in particular, may suggest that groups of species coflower more strongly (clustering) with each other than with other species (over-dispersion) in the community. Such a finding would indicate that competitive and facilitative interactions do not act alone but instead act simultaneously to mediate the assembly of coflowering communities. We illustrate our conceptual framework in four diverse coflowering communities in the serpentine seeps in northern California. Pivotal results. Our coflowering networks vary in size and degree but not in overall connectance, suggesting that both intrinsic community features (species richness) and ecological constraints (length of flowering season) play a role in mediating coflowering community structure (distribution of frequency and intensity of flowering overlap among plant species). We show, for the first time, that groups of species tend to coflower more strongly with each other than with other species in a community, supporting the idea that competition and facilitation are not mutually exclusive processes mediating coflowering community assembly. Our results show that the degree of modularity is not sensitive to the number of coflowering species within each community, suggesting that ecological factors may be more important in driving this pattern. Conclusions. Coflowering networks have the potential to advance our understanding of the causes and consequences of flowering overlap in diverse plant communities by revealing a more in-depth and novel characterization of coflowering community structure. Such characterization will allow for a better understanding of the importance of coflowering patterns in mediating the structure of plant-pollinator interactions.

Premise of the Study: There is growing interest in understanding plant–plant interactions via pollen transfer at the community level. Studies on the structure and spatial variability of pollen transfer networks have been valuable to this understanding. However, there is high variability in the intensity of sampling used to characterize pollen transfer interactions, which could influence network structure. To date, there is no knowledge of how sampling effort influences the richness of pollen on stigmas and thereby transfer interactions observed, nor how this may vary across species and study sites. Methods: We use rarefaction curves on 16 species to characterize the relationship between sampling effort (number of stigmas analyzed) and the richness of pollen transfer interactions recorded. We further assess variability in this relationship among species, plant community types, and sites within a single plant community. Key Results: We show high among-species variation in the amount of sampling required to sufficiently characterize interspecific pollen transfer. We further reveal variability in the sampling effort-interaction richness relationship among different plant communities and even for the same species growing in different sites. Conclusions: The wide heterogeneity in the sampling effort required to accurately characterize pollen transfer interactions observed has the potential to influence the characterization of pollen transfer dynamics. Thus, sampling completeness should be considered in future studies to avoid overestimation of modularity and specialization in pollen transfer networks that may bias the predicted causes and expected consequences of such processes for plant–plant interactions.

Background: Pollen transfer via animals is necessary for reproduction by ~80 % of flowering plants, and most of these plants live in multispecies communities where they can share pollinators. While diffuse plant-pollinator interactions are increasingly recognized as the rule rather than the exception, their fitness consequences cannot be deduced from flower visitation alone, so other proxies, functionally closer to seed production and amenable for use in a broad variety of diverse communities, are necessary. Scope: We conceptually summarize how the study of pollen on stigmas of spent flowers can reflect key drivers and functional aspects of the plant-pollinator interaction (e.g. competition, facilitation or commensalism). We critically evaluate how variable visitation rates and other factors (pollinator pool and floral avoidance) can give rise to different relationships between heterospecific pollen and (1) conspecific pollen on the stigma and (2) conspecific tubes/grain in the style, revealing the complexity of potential interpretations. We advise on best practices for using these proxies, noting the assumptions and caveats involved in their use, and explicate what additional data are required to verify interpretation of given patterns. Conclusions: We conclude that characterizing pollen on stigmas of spent flowers provides an attainable indirect measure of pollination interactions, but given the complex processes of pollen transfer that generate patterns of conspecific-heterospecific pollen on stigmas these cannot alone determine whether competition or facilitation are the underlying drivers. Thus, functional tests are also needed to validate these hypotheses.