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Understanding the functional consequences of biodiversity loss is a major goal of ecology. Animal-mediated pollination is an essential ecosystem function and service provided to mankind. However, little is known how pollinator diversity could affect pollination services. Using a substitutive design, we experimentally manipulated functional group (FG) and species richness of pollinator communities to investigate their consequences on the reproductive success of an obligate out-crossing model plant species, Raphanus sativus. Both fruit and seed set increased with pollinator FG richness. Furthermore, seed set increased with species richness in pollinator communities composed of a single FG. However, in multiple-FG communities, highest species richness resulted in slightly reduced pollination services compared with intermediate species richness. Our analysis indicates that the presence of social bees, which showed roughly four times higher visitation rates than solitary bees or hoverflies, was an important factor contributing to the positive pollinator diversity–pollination service relationship, in particular, for fruit set. Visitation rate at different daytimes, and less so among flower heights, varied among social bees, solitary bees and hoverflies, indicating a niche complementarity among these pollinator groups. Our study demonstrates enhanced pollination services of diverse pollinator communities at the plant population level and suggests that both the niche complementarity and the presence of specific taxa in a pollinator community drive this positive relationship.

Secondary metabolites that are present in floral nectar have been hypothesized to enhance specificity in plant–pollinator mutualism by reducing larceny by non‐pollinators, including microorganisms that colonize nectar. However, few studies have tested this hypothesis. Using synthetic nectar, we conducted laboratory and field experiments to examine the effects of five chemical compounds found in nectar on the growth and metabolism of nectar‐colonizing yeasts and bacteria, and the interactive effects of these compounds and nectar microbes on the consumption of nectar by pollinators. In most cases, focal compounds inhibited microbial growth, but the extent of these effects depended on compound identity, concentration, and microbial species. Moreover, most compounds did not substantially decrease sugar metabolism by microbes, and microbes reduced the concentration of some compounds in nectar. Using artificial flowers in the field, we also found that the common nectar yeast Metschnikowia reukaufii altered nectar consumption by small floral visitors, but only in nectar containing catalpol. This effect was likely mediated by a mechanism independent of catalpol metabolism. Despite strong compound‐specific effects on microbial growth, our results suggest that the secondary metabolites tested here are unlikely to be an effective general defense mechanism for preserving nectar sugars for pollinators. Instead, our results indicate that microbial colonization of nectar could reduce the concentration of secondary compounds in nectar and, in some cases, reduce deterrence to pollinators.

Summary Floral colour mediates plant–pollinator interactions by often signalling floral resources. In this sense, hummingbird‐pollinated flowers are frequently red‐coloured, and there are two tentative hypotheses to explain this pattern: 1. hummingbirds are attracted to red due its easier detection and 2. bees are sensorially excluded from red flowers. The second hypothesis is based on bees’ red colour blindness, which lead them to be less frequent and less important than hummingbirds as pollinators of red‐reflecting flowers. Here, we untangled the role of different flower traits mediating plant–pollinator interactions and empirically tested the above hypotheses. We chose Costus arabicus due to its synchronopatric white‐ and pink‐flowered individuals and its bee and hummingbird pollination system. Although pink flowers are not totally achromatic as pure red ones, they show an achromaticity degree that could drive bee exclusion. Specifically, we tested whether differences on red reflectance work attracting hummingbirds or excluding bees and the consequent implications for the plant's reproduction. Flower colour morphs of C. arabicus do differ neither in morphology nor in nectar sugar content. Moreover, white and pink flowers can be discriminated by the bees’ and hummingbirds’ colour vision system. Both groups are able to discriminate the red colour variation morph on the flower petals, the white flowers being more easily detected by bees and the pink flowers by hummingbirds. Bees preferentially visited the white flowers, whereas hummingbirds visited both colours at the same rate – both patterns corroborating the second hypothesis. Pollen loads deposited on stigmas did not differ between flower colour morphs, indicating that bees and hummingbirds play a similar role in the overall pollen deposition. However, bees are more likely to self‐pollinate than hummingbirds. Self‐pollination limits C. arabicus reproduction, and red‐reflecting flowers may be better pollinated by discouraging bee visitation. Therefore, the intraspecific colour variation is driving flowers to show colour‐related different levels of generalization. Our results support the ‘bee avoidance’ rather than the ‘hummingbird preference’ hypothesis. Sensory exclusion of bees seems to be the pressure for red‐reflecting flowers evolution, driving specialization in hummingbird‐pollinated flowers due to the costs of bee pollination on plant reproduction. Lay Summary

A fundamental goal of ecology is to predict how strongly one species affects the abundance of another. However, our ability to do so is hindered by the fact that interaction outcomes are notoriously variable in space and time (i.e. context‐dependent) and we lack a predictive understanding of the factors that drive this context‐dependence. Determining whether abiotic factors, in particular, predictably shift the outcome of species interactions is of critical importance for many contemporary problems, from forecasting climate change impacts to predicting the efficacy of weed biocontrol. In this essay, we highlight the context‐dependent nature of interactions between plants and their pollinators and herbivores. We advocate for approaches that will identify whether particular abiotic factors predictably shift how strongly these interactions influence plant abundance and/or population growth. We review long‐standing theory that describes how abiotic context should influence the selective impacts of pollinators and herbivores on plants and articulate why this theory requires modification to predict population‐level effects. Finally, we propose several empirical approaches to address gaps in existing knowledge: (i) experiments across broad abiotic gradients to determine whether the outcome of interactions between pollinators or herbivores and plants varies consistently with changing abiotic conditions; (ii) experiments that manipulate the underlying environmental gradient to elucidate whether the abiotic factor that correlates with interaction outcome is causal; and (iii) seed addition studies to explore how strongly seedling recruitment correlates with seed input (as affected by pollen limitation or herbivory) and to quantify how the strength of the seed‐to‐seedling linkage is influenced by the underlying abiotic gradient. Synthesis. Our understanding of the underlying drivers of context‐dependent plant–animal interactions is currently not well developed. Progress in this area is essential to better predict when and where species interactions will alter the responses of plant populations to environmental changes as well as to develop more robust theory. Experiments aimed at explicitly exploring the role of abiotic factors in mediating the population‐level impact of pollen limitation and herbivory could determine the extent to which variation in the abiotic environment predictably shifts the outcome of these interactions.

Premise of the study: To study pollination networks in a changing environment, we need accurate, high-throughput methods. Previous studies have shown that more highly resolved networks can be constructed by studying pollen loads taken from bees, relative to field observations. DNA metabarcoding potentially allows for faster and finer-scale taxonomic resolution of pollen compared to traditional approaches (e.g., light microscopy), but has not been applied to pollination networks. Methods: We sampled pollen from 38 bee species collected in Florida from sites differing in forest management. We isolated DNA from pollen mixtures and sequenced rbcL and ITS2 gene regions from all mixtures in a single run on the Illumina MiSeq platform. We identified species from sequence data using comprehensive rbcL and ITS2 databases. Results: We successfully built a proof-of-concept quantitative pollination network using pollen metabarcoding. Discussion: Our work underscores that pollen metabarcoding is not quantitative but that quantitative networks can be constructed based on the number of interacting individuals. Due to the frequency of contamination and false positive reads, isolation and PCR negative controls should be used in every reaction. DNA metabarcoding has advantages in efficiency and resolution over microscopic identification of pollen, and we expect that it will have broad utility for future studies of plant–pollinator interactions.

The role of pollination in the success of invasive plants needs to be understood because invasives have substantial effects on species interactions and ecosystem functions. Previous research has shown both that reproduction of invasive plants is often pollen limited and that invasive plants can have high seed production, motivating the questions: How do invasive populations maintain reproductive success in spite of pollen limitation? What species traits moderate pollen limitation for invaders? We conducted a phylogenetic meta-analysis with 68 invasive, 50 introduced noninvasive and 1931 native plant populations, across 1249 species. We found that invasive populations with generalist pollination or pollinator dependence were less pollen limited than natives, but invasives and introduced noninvasives did not differ. Invasive species produced 3× fewer ovules/flower and > 250× more flowers per plant, compared with their native relatives. While these traits were negatively correlated, consistent with a tradeoff, this did not differ with invasion status. Invasive plants that produce many flowers and have floral generalisation are able to compensate for or avoid pollen limitation, potentially helping to explain the invaders’ reproductive successes.

Community studies have shown that plant species are often pollinated by multiple pollinators; however, networks of heterospecific pollen transfer (HPT) in natural communities remain largely unexplored. We analyzed pollen deposition on stigmas of 57 flowering species to build a picture of plant-plant interactions via HPT in a biodiverse alpine meadow in southwest China. Plant species were categorized as pollen donors or recipients by their link numbers and link qualities. We identified 3609 heterospecific pollen grains, representing 410 links among 69 pollen species. Each plant species received on average 7.2 pollen species and donated its pollen to 5.5 species; only a few species donated or received large amounts of pollen or pollen from a large number of species. Compared to specialized plants, generalized plants tended to receive more heterospecific pollen but exported no more pollen to other species. Plant position in the network was related to both floral traits (stigma position) and pollinator generalization level. When different species share the same pollinator, bidirectional HPT may occur, but this was rarely observed in the species-rich community, indicating that interspecific pollen interference was largely unidirectional. Our study highlights the importance of understanding how sympatric flowering plants reduce deleterious effects of HPT, for example via stigma position. This study is the first to present a pollen transfer network for an entire community and to unravel its properties using directed network analysis.

Both leaves and petals are covered in a cuticle, which itself contains and is covered by cuticular waxes. The waxes perform various roles in plants’ lives, and the cuticular composition of leaves has received much attention. To date, the cuticular composition of petals has been largely ignored. Being the outermost boundary between the plant and the environment, the cuticle is the first point of contact between a flower and a pollinator, yet we know little about how plant–pollinator interactions shape its chemical composition. Here, we investigate the general structure and composition of floral cuticular waxes by analysing the cuticular composition of leaves and petals of 49 plant species, representing 19 orders and 27 families. We show that the flowers of plants from across the phylogenetic range are nearly devoid of wax crystals and that the total wax load of leaves in 90% of the species is higher than that of petals. The proportion of alkanes is higher, and the chain lengths of the aliphatic compounds are shorter in petals than in leaves. We argue these differences are a result of adaptation to the different roles leaves and petals play in plant biology.

Flowers act as multisensory billboards to pollinators by using a range of sensory modalities such as visual patterns and scents. Different floral organs release differing compositions and quantities of the volatiles contributing to floral scent, suggesting that scent may be patterned within flowers. Early experiments suggested that pollinators can distinguish between the scents of differing floral regions, but little is known about how these potential scent patterns might influence pollinators. We show that bumblebees can learn different spatial patterns of the same scent, and that they are better at learning to distinguish between flowers when the scent pattern corresponds to a matching visual pattern. Surprisingly, once bees have learnt the spatial arrangement of a scent pattern, they subsequently prefer to visit novel unscented flowers that have an identical arrangement of visual marks, suggesting that multimodal floral signals may exploit the mechanisms by which learnt information is stored by the bee.

Predicting how communities re-arrange in response to changes in species composition remains a key challenge in ecology. Migratory species, which enter and leave communities across latitudinal gradients, offer us a unique opportunity to evaluate community- and species-level responses to a shift in community composition. We focused on a migratory hummingbird and the communities that host it along a latitudinal and species diversity gradient. Our results show higher niche overlap in more diverse communities, allowing resident species to compensate for the loss of the migrant in providing pollination services. Contrastingly, in less diverse communities, the migrant behaves as a specialist, monopolizing abundant resources. In its absence, its role is not fully covered by resident species, resulting in a decrease in the fruit set of the migrant's preferred plant species. These results help us understand the potential impacts of biodiversity loss and have important implications for community persistence given expected changes in the migratory behaviours of some species. © 2020 The Author(s).