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Floral traits are hypothesized to evolve primarily in response to selection by pollinators. However, selection can also be mediated by other environmental factors. To understand the relative importance of pollinator-mediated selection and its variation among trait and pollinator types, we analyzed directional selection gradients on floral traits from experiments that manipulated the environment to identify agents of selection. Pollinator-mediated selection was stronger than selection by other biotic factors (e.g., herbivores), but similar in strength to selection by abiotic factors (e.g., soil water), providing partial support for the hypothesis that floral traits evolve primarily in response to pollinators. Pollinator-mediated selection was stronger on pollination efficiency traits than on other trait types, as expected if efficiency traits affect fitness via interactions with pollinators, but other trait types also affect fitness via other environmental factors. In addition to varying among trait types, pollinator-mediated selection varied among pollinator taxa: selection was stronger when bees, long-tongued flies, or birds were the primary visitors than when the primary visitors were Lepidoptera or multiple animal taxa. Finally, reducing pollinator access to flowers had a relatively small effect on selection on floral traits, suggesting that anthropogenic declines in pollinator populations would initially have modest effects on floral evolution.

Wild and managed bees are well documented as effective pollinators of global crops of economic importance. However, the contributions by pollinators other than bees have been little explored despite their potential to contribute to crop production and stability in the face of environmental change. Non-bee pollinators include flies, beetles, moths, butterflies, wasps, ants, birds, and bats, among others. Here we focus on non-bee insects and synthesize 39 field studies from five continents that directly measured the crop pollination services provided by non-bees, honey bees, and other bees to compare the relative contributions of these taxa. Non-bees performed 25–50% of the total number of flower visits. Although non-bees were less effective pollinators than bees per flower visit, they made more visits; thus these two factors compensated for each other, resulting in pollination services rendered by non-bees that were similar to those provided by bees. In the subset of studies that measured fruit set, fruit set increased with non-bee insect visits independently of bee visitation rates, indicating that non-bee insects provide a unique benefit that is not provided by bees. We also show that non-bee insects are not as reliant as bees on the presence of remnant natural or seminatural habitat in the surrounding landscape. These results strongly suggest that non-bee insect pollinators play a significant role in global crop production and respond differently than bees to landscape structure, probably making their crop pollination services more robust to changes in land use. Non-bee insects provide a valuable service and provide potential insurance against bee population declines.

The evolution of floral traits in animal-pollinated plants involves the interaction between flowers as signal senders and pollinators as signal receivers. Flower colors are very diverse, effect pollinator attraction and flower foraging behavior, and are hypothesized to be shaped through pollinator-mediated selection. However, most of our current understanding of flower color evolution arises from variation between discrete color morphs and completed color shifts accompanying pollinator shifts, while evidence for pollinator-mediated selection on continuous variation in flower colors within populations is still scarce. In this review, we summarize experiments quantifying selection on continuous flower color variation in natural plant populations in the context of pollinator interactions. We found that evidence for significant pollinator-mediated selection is surprisingly limited among existing studies. We propose several possible explanations related to the complexity in the interaction between the colors of flowers and the sensory and cognitive abilities of pollinators as well as pollinator behavioral responses, on the one hand, and the distribution of variation in color phenotypes and fitness, on the other hand. We emphasize currently persisting weaknesses in experimental procedures, and provide some suggestions for how to improve methodology. In conclusion, we encourage future research to bring together plant and animal scientists to jointly forward our understanding of the mechanisms and circumstances of pollinator-mediated selection on flower color.

Conserving and restoring semi-natural habitat, i.e. enhancing landscape complexity, is one of the main strategies to mitigate pollinator decline in agricultural landscapes. However, we still have limited understanding of how landscape complexity shapes pollinator communities in both crop and non-crop habitat, and whether pollinator responses to landscape complexity vary with their association with mass-flowering crops. Here, we surveyed pollinator communities on mass-flowering leek crops and in nearby semi-natural habitat in landscapes of varying complexity. Surveys were done before and during crop bloom and distinguished between pollinators that visit the crop frequently (dominant), occasionally (opportunistic), or not at all (non-crop). Forty-seven per cent of the species in the wider landscape were also observed on leek flowers. Crop pollinator richness increased with local pollinator community size and increasing landscape complexity, but relationships were stronger for opportunistic than for dominant crop pollinators. Relationships between pollinator richness in semi-natural habitats and landscape complexity differed between groups with the most pronounced positive effects on non-crop pollinators. Our results indicate that while dominant crop pollinators are core components of crop pollinator communities in all agricultural landscapes, opportunistic crop pollinators largely determine species-richness responses and complex landscapes are local hotspots for both biodiversity conservation and potential ecosystem service provision.

Abstract Background The male fitness pathway, from pollen production to ovule fertilization, is thought to strongly influence reproductive trait evolution in animal-pollinated plants. This pathway is characterized by multiple avenues of pollen loss which may lead to reductions in male fitness. However, empirical data on the mechanistic processes leading to pollen loss during transport are limited, and we therefore lack a comprehensive understanding of how male fitness is influenced by each step in the pollination process. Scope This review assesses the history of studying male function in plants and identifies critical gaps in our understanding of the ecology and evolution of pollen transport. We explore male reproductive function along the steps of the pathway to paternity and discuss evolutionary options to overcome barriers to siring success. In particular, we present a newly emerging idea that bodies of pollinators function as a dynamic arena facilitating intense male–male competition, where pollen of rival males is constantly covered or displaced by competitors. This perspective extends the pollen-competitive arena beyond the confines of the stigma and style, and highlights the opportunity for important new breakthroughs in the study of male reproductive strategies and floral evolution.

Current notions of \"pollinator decline\" and \"pollination crisis\" mainly arose from studies on pollinators of economic value in anthropogenic ecosystems of mid-latitude temperate regions. Comprehensive long-term pollinator data from biologically diverse, undisturbed communities are needed to evaluate the actual extent of the so-called \"global pollination crisis.\" This paper analyzes the long-term dynamics of pollinator abundance in undisturbed Mediterranean montane habitats using pollinator visitation data for 65 plant species collected over two decades. Objectives are (1) to elucidate patterns of long-term changes in pollinator abundance from the perspectives of individual plant species, major pollinator groups, and the whole plant community and (2) to propose a novel methodological implementation based on combining a planned missing data design with the analytical strength of mixed effects models, which allows one to draw community-wide inferences on long-term pollinator trends in species-rich natural habitats. Probabilistic measurements (\"patch visitation probability\" and \"flower visitation probability\" per time unit) were used to assess pollinator functional abundance for each plant species on two separate, randomly chosen years. A total of 13,054 pollinator censuses accounting for a total watching effort of 2,877,039 flower-min were carried out on 299 different dates. Supra-annual unstability in pollinator functional abundance was the rule, with visitation probability to flowering patches and/or individual flowers exhibiting significant heterogeneity between years in the majority of plant species (83%). At the plantcommunity level, there was a significant linear increase in pollinator functional abundance over the study period. Probability of pollinator visitation to flowering patches and individual flowers increased due to increasing visitation by small solitary bees and, to a lesser extent, small beetles. Visitation to different plant species exhibited contrasting changes, and insect orders and genera differed widely in sign and magnitude of linear abundance trends, thus exemplifying the complex dynamics of community-wide changes in pollinator functional abundance. Results of this investigation indicate that pollinator declines are not universal beyond anthropogenic ecosystems; stress the need for considering broader ecological scenarios and comprehensive samples of plants and pollinators; and illustrate the crucial importance of combining ambitious sampling designs with powerful analytical schemes to draw reliable inferences on pollinator trends at the plant community level.

Pollinator service is essential for successful sexual reproduction and long-term population persistence of animal-pollinated plants, and innumerable studies have shown that insufficient service by pollinators results in impaired sexual reproduction (\"pollen limitation\"). Studies directly addressing the predictors of variation in pollinator service across species or habitats remain comparatively scarce, which limits our understanding of the primary causes of natural variation in pollen limitation. This paper evaluates the importance of pollination-related features, evolutionary history, and environment as predictors of pollinator service in a large sample of plant species from undisturbed montane habitats in southeastern Spain. Quantitative data on pollinator visitation were obtained for 191 insect-pollinated species belonging to 142 genera in 43 families, and the predictive values of simple floral traits (perianth type, class of pollinator visitation unit, and visitation unit dry mass), phylogeny, and habitat type were assessed. A total of 24,866 pollinator censuses accounting for 5,414,856 flower-minutes of observation were conducted on 510 different dates. Flowering patch and single flower visitation probabilities by all pollinators combined were significantly predicted by the combined effects of perianth type (open vs. restricted), class of visitation unit (single flower vs. flower packet), mass of visitation unit, phylogenetic relationships, and habitat type. Pollinator composition at insect order level varied extensively among plant species, largely reflecting the contrasting visitation responses of Coleoptera, Diptera, Hymenoptera, and Lepidoptera to variation in floral traits. Pollinator composition had a strong phylogenetic component, and the distribution of phylogenetic autocorrelation hotspots of visitation rates across the plant phylogeny differed widely among insect orders. Habitat type was a key predictor of pollinator composition, as major insect orders exhibited decoupled variation across habitat types in visitation rates. Comprehensive pollinator sampling of a regional plant community has shown that pollinator visitation and composition can be parsimoniously predicted by a combination of simple floral features, habitat type, and evolutionary history. Ambitious community-level studies can help to formulate novel hypotheses and questions, shed fresh light on long-standing controversies in pollination research (e.g., \"pollination syndromes\"), and identify methodological cautions that should be considered in pollination community studies dealing with small, phylogenetically biased plant species samples.

Aim Understanding fire effects on pollinators is critical in the context of fire regime changes and the global pollination crisis. Through a systematic and quantitative review of the literature, we provide the first global assessment of pollinator responses to fire. We hypothesize that pollinators increase after fire and during the early postfire succession stages; however, high fire frequency has the opposite effect, decreasing pollinators. Location Terrestrial ecosystems, excluding Antarctica. Time period Data collected from 1973 to 2017. Major taxa studied Insects (Coleoptera, Diptera, Hymenoptera and Lepidoptera) and a few bird species. Methods We first compiled available studies across the globe that assessed fire effects on pollinator communities. Then, by means of hierarchical meta‐analyses, we evaluated how different fire regime parameters (fire frequency, postfire time and fire type) and habitat characteristics affect the abundance and richness of animals that act as pollinators. We also explored to what extent the responses vary among taxa groups and life history traits of pollinators (sociality system, nest location and feeding specialization), and among biomes. Results The overall effect size of fire on pollinator abundance and richness across all studies was positive. Fire effect was especially clear and significant in early postfire communities, after wildfires, and for Hymenoptera. Taxonomic resolution influenced fire effects, where only studies at the species/genus and family levels showed significant effects. The main exceptions were recurrent fires that showed a negative effect, and especially wildfire effects on Lepidoptera abundance that showed a significant negative response. Main conclusions Pollinators tend to be promoted after a wildfire event. However, short fire intervals may threat pollinators, and especially lepidopterans. Given the current fire regime changes at the global scale, it is imperative to monitor postfire pollinators across many ecosystems, as our results suggest that fire regime is critical in determining the dynamics of pollinator communities.