Functional traits, particularly those that impact fitness, can shape the ecological and evolutionary relationships among coexisting species of the same trophic level. Thus, examining these traits and properties of their distributions (underdispersion, overdispersion) within communities can provide insights into key ecological interactions (e.g., competition, facilitation) involved in community assembly. For instance, the distribution of floral colors in a community may reflect pollinator-mediated interactions between sympatric plant species, and the phylogenetic distribution of color can inform how evolutionary contingencies can continue to shape extant community assemblages. Additionally, the abundance and species richness of the local habitat may influence the type or strength of ecological interactions among co-occurring species. To evaluate the impact of community size and species richness on mechanisms shaping the distribution of ecologically relevant traits, we examined how floral color (defined by pollinator color vision models) is distributed within co-flowering assemblages. We modeled floral reflectance spectra of 55 co-flowering species using honeybee (Apis mellifera) and syrphid fly (Eristalis tenax) visual systems to assess the distributions of flower color across 14 serpentine seep communities in California. We found that phylogenetic relatedness had little impact on the observed color assemblages. However, smaller seep communities with lower species richness were more overdispersed for flower color than larger, more species-rich communities. Results support that competitive exclusion could be a dominant process shaping the species richness of flower color in smaller-sized communities with lower species richness, but this is less detectable or overwhelmed by other processes at larger, more speciose communities.

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.

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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.

Ecological interactions within a community shape the structure of ecosystems and influence ecosystem function. Plant-pollinator interactions exist as mutualistic exchange networks that may collapse as habitat loss occurs, thereby threatening the overall health of an ecosystem. Understanding the impacts of human-mediated habitat disturbance on ecological interactions is therefore crucial for conservation efforts. Archbold Biological Station (ABS) in Venus, Florida contains over 2000 hectares of protected Florida scrub habitat nested within a human-dominated environment that is threatened by anthropogenic habitat disturbance. In past studies, over 113 bee species and 157 associated host plants, many endemic to the Lake Wales Ridge, have been found on ABS property, providing an understanding of this system's plant-bee network. Using those data as a baseline, this study investigated the effects of varying levels of mechanical habitat disturbance intensity on the diversity and network structure of plant-bee interaction networks. Flowering plant abundance, richness, diversity, and composition as well as bee abundance and composition were significantly different across mechanical habitat disturbance levels. Interactions between bees and flowering plants also differed with varying disturbance intensity. From these results, it is clear that plants, bees and interactions between them are impacted by mechanical habitat disturbance in this system. This project informs management efforts not only for natural systems with the threat of alteration, but also for agricultural systems, many of which heavily rely on flower visitation by bee pollinators. This research also contributes to the growing field of interaction ecology by increasing understanding of habitat alteration effects on a valuable ecological interaction and ultimately ecosystem function.

Doctor of Philosophy / Division of Biology / Anthony Joern / John Blair / Ecological communities are comprised both of species and their interactions. The importance of species interactions is embraced by ecological network analysis, a framework used to identify non-random patterns in species interactions, and the consequences of these patterns for maintaining species diversity. Here, I investigated environmental drivers of the structure of plant-pollinator and plant-herbivore networks. Specifically, I asked: (1) Do global-scale climate gradients shape mutualistic and antagonistic networks? (2) At a landscape scale (within a 3,487 ha research site), how do contrasting regimes of major grassland disturbances - fire frequency and grazing by bison (Bison bison) - shape plant-pollinator network structure? (3) How do fire and grazing affect plant-grasshopper network structure? And, (4) What is the role of plant species diversity in determining plant-herbivore network structure? At the global scale, variability in temperature was the key climatic factor regulating both antagonistic and mutualistic network structural properties. At the landscape scale, fire and grazing had major consequences for plant-pollinator and plant-herbivore communities. In particular, bison grazing increased network complexity and resistance to species loss for both plant-pollinator and plant-herbivore systems. Results from an experimental grassland restoration that manipulated plant diversity suggest that plant diversity directly affects plant-herbivore structure and increases network stability. Collectively, these results suggest that environmental gradients and plant species diversity regulate the network structure of ecological communities. Determining how the structure of ecological interactions change with environmental conditions and species diversity improves our ability to identify vulnerable communities, and to predict responses of biodiversity to global change.

The study of foraging behaviour in plant-pollinator mutualisms has benefitted from the use of artificial flowers to manipulate floral display traits and the delivery of floral rewards. The two most common floral rewards are pollen and nectar; some pollinators, such as bees, are obliged to collect both for survival and reproduction. While flexible designs for artificial flowers providing nectar rewards abound, useful designs for artificial flowers that dispense pollen are few. This disparity mirrors a heavy emphasis on nectar collection in the study of pollinator foraging behaviour. In this study we describe a novel, easily constructed and modifiable artificial flower that dispenses flexible amounts of pollen via an ‘anther’ composed of a chenille stem. Using controlled lab assays, we show that more pulverized honeybee pollen is collected by bumblebee (Bombus impatiens) workers at chenille stem feeders than at dish-type feeders. We suggest that the paucity of studies examining pollinator cognition in the context of pollen rewards might be partly remedied if researchers had access to inexpensive and easily adjustable pollen-offering surrogate flowers.

Bees provide a vital pollination service to many important crops and wildflowers yet are experiencing population declines across European and North American agricultural landscapes. The conservation of bee communities on farmland is a priority of the UK agri-environment schemes, which support pollinators through the provision of natural and semi-natural habitat, foraging resources and nesting sites. Data are needed to evaluate the effectiveness of these interventions in supporting bee communities and to allow the refinement of effective, evidence-based policy. This thesis examines the effectiveness of agri-environment management, though the provision of natural habitat, foraging resources and nesting sites, in supporting solitary bees, an important group of pollinators of which there are 250 species in the UK. Cavity nesting solitary bees and wasps were surveyed on 19 farms situated across central southern England, ranging from farms under no agri-environment scheme, to farms showcasing higher-level agri-environment management. Data on bee and wasp communities was collected by deploying solitary bee nest boxes. These nest boxes are marketed widely as nesting resources for solitary bees and are provided to farmers as part of Higher Level agri-environment schemes. Over the course of the study, 4002 solitary bees and wasps, comprising 10 species, were recorded. Natural Habitat: A positive relationship was identified between the abundance and species richness of bees and wasps and the proportion of natural habitat across farms. The availability of natural habitat also had a positive influence on the structural stability of bee and wasp parasitism networks; a positive association was identified with network link density. Link density measures the mean number of links per species within a network. Higher measures of link density are believed to confer greater resilience to species loss as individuals have more flexibility to switch interaction partner, limiting the risk of a cascade of secondary species extinctions. This set of results is encouraging, suggesting that the natural habitat types being promoted are effective in supporting solitary bee and wasp communities. Foraging resources: Examination of bee foraging preferences, through next generation sequencing of brood cell pollen DNA, demonstrated that the agri-environment scheme sown wildflower mixes do not support the foraging requirements of solitary bees effectively. Of the 15 plants included currently in the wildflower mixes that were recorded as present on the study farms, pollen from only one species, Ranunculus acris, was used by the bees. Rosa canina was identified as the most popular forage plant. The leaves of this species are also a preferred nesting material for Megachile leafcutter bees, providing strong justification for the inclusion of R. canina within the selection of hedgerow plants encouraged by agri-environment schemes. Tripleurospermum inodorum and Trifolium repens were also identified as good candidates for inclusion in wildflower seed mixtures. Nesting sites: A strong positive relationship between the density of solitary bee nest boxes and the rate of brood cell parasitism was identified, indicating that a high local density of nest boxes may expose bee larvae to a higher risk of parasitism. An enhanced risk of larval mortality could counteract the benefit of additional nest site provision. No significant effect of nest box provision on nest box colonization was identified across these study sites, suggesting that their placement across landscapes to encourage more pollinators may be counter-productive. It would be prudent to advise, given the results of this study, for the provision of a small number of dispersed nest boxes, this might more accurately mimic the availability of nesting resources in nature and reduce the risk of enhanced parasitism rates.

The overall aim of the study was to assess the risk of gene flow from Brassica crops by insectmediated pollen transport. I measured the viability of pollen in Brassica flowers throughout crop development and compared this with the viability of pollen transported by insects inside and outside one early- and one late-season crop. In order to evaluate the relative importance of different species in pollen transport, I measured abundance of flower visitors during crop development, and measured the foraging behaviour of five key pollinator species throughout the growing season, in relation to variation in microclimate, crop phenology and the relative abundance of other pollinator species competing for flower resources. Flower visiting insects of Brassica rapa crops were highly diverse, and their abundance and diversity changed with crop phenology. I found similar abundances at the family level for both crops studied, although capture rates were greater in the early- than in the late-season crop. Across flowering development, the greatest numbers of insects were captured at the peak of flowering for both crops. During the flowering period, Diptera was the most abundant order collected, followed by Hymenoptera. The most abundant family in Hymenoptera was Apidae which tracked crop development in both fields, with greater numbers of insects captured inside than outside the field. Standardized-count pollen loads were smaller in Diptera than in Hymenoptera. Of the five key pollinator species sampled, Lasioglossum sordidum (Hymenoptera: Halictidae), Apis mellifera (Hymenoptera: Apidae) and Bombus terrestris (Hymenoptera: Apidae) transported similar pollen loads, which were much greater than those carried by Eristalis tenax (Diptera: Syrphidae) and Melangyna novae-zealandiae (Diptera: Syrphidae). The numbers of insects captured outside of the crop were 10% and 33% of the totals captured inside for the early- and the late-season crop, respectively. The proportion of insects entering versus leaving the crop varied considerably across species, crops and trap location (i.e., whether traps were inside or 50 m outside the border of the crop). However, it is worth noting that not uncommonly more insects were attracted into the crop early in the season, staying there rather than leaving, and then when flowers started to disappear there was a massive escape of insects leaving. This research provides evidence for the influence of crop age on the foraging behaviour of key pollinators and for species-specific variation in the foraging behaviour of Brassica visitors with crop development. Temporal variation in the rate and variability of movement between flowers, and the duration and variability in time spent on each flower, throughout the growing season differed markedly between pollinator species. Flower density, plant density, and the abundance of other insects contributed to the observed variation in pollinator behavioural activity for A. mellifera, E. tenax, M. novae-zelandiae and L. sordidum. Bombus terrestris had the greatest rates and variability of movement, and the greatest rates of flower visitation among all key pollinators studied. Therefore B. terrestris might contribute to gene flow to a greater extent than other key pollinators. Additionally B. terrestris had the greatest variability in the rate of movement, increasing the risk of pollen movement over long distances. In summary, I found that (i) insect abundance and diversity changed with crop phenology and Diptera was the most abundant order collected, (ii) flower density, plant density, and the abundance of other insect pollinators were important factors explaining pollinator behaviour for all key pollinators, except B. terrestris, (iii) B. terrestris might contribute to gene flow to a greater extent than other key pollinators, because it has a greater rate of flower visitation and a greater flight distance between flowers than other pollinators, and (iv) pollen viability tended to decrease with crop development and declined sharply even just 50 m outside the edge of the crop.