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The responses of animal pollinators to the spatially heterogeneous distribution of floral resources are important for plant reproduction, especially in species-rich plant communities. We explore how responses of pollinators to floral resources varied across multiple spatial scales and studied the responses of two nectarivorous bird species (Cape sugarbird Promerops cafer, orange-breasted sunbird Anthobaphes violacea) to resource distributions provided by communities of co-flowering Protea species (Proteaceae) in South African fynbos. We used highly resolved maps of about 125 000 Protea plants at 27 sites and estimated the seasonal dynamics of standing crop of nectar sugar for each plant to describe the spatiotemporal distribution of floral resources. We recorded avian population sizes and the rates of bird visits to > 1300 focal plants to assess the responses of nectarivorous birds to floral resources at different spatial scales. The population sizes of the two bird species responded positively to the amount of sugar resources at the site scale. Within sites, the effects of floral resources on pollinator visits to plants varied across scales and depended on the resources provided by individual plants. At large scales (radii > 25 m around focal plants), high sugar density decreased per-plant visitation rates, i.e. plants competed for animal pollinators. At small scales (radii < 5 m around focal plants), we observed either competition or facilitation for pollinators between plants, depending on the sugar amount offered by individual focal plants. In plants with copious sugar, per-plant visitation rates increased with increasing local sugar density, but visitation rates decreased in plants with little sugar. Our study underlines the importance of scale-dependent responses of pollinators to floral resources and reveals that pollinators' responses depend on the interplay between individual floral resources and local resource neighbourhood. [ABSTRACT FROM AUTHOR]

Pollination syndromes predict the principal animal pollinators of plant species from floral traits. However, the existence of discrete pollination syndromes and the relevance of these syndromes for the effects of animal pollinators on plant reproduction are controversial and a quantitative, trait-based approach may refine the concept of pollination syndromes. Thus far, however, very few studies have simultaneously analysed the importance of signalling, accessibility and reward traits of flowers for the effects of animal pollinators on plant reproduction of multiple species. The genus Protea is an ideal study system because the members of this genus exhibit remarkable variation in functional traits and Protea species are visited by several guilds of animal pollinators, such as birds and insects. We experimentally investigated seven Protea species from the South African fynbos to assess variability in reproduction (seed set, seed mass, germination) in response to pollinator exclusion (full-exclusion, bird-exclusion) and hand-pollination (self-pollen, cross-pollen). To generalise the findings of the experiment, we conducted a meta-analysis of the effects of bird-exclusion on seed set of 14 Protea species, associating effect sizes of bird-pollination with signalling (inflorescence length and diameter), accessibility (inflorescence and style length) and reward traits (length of pollen presenter, nectar volume and sugar concentration). All Protea species were self-compatible, but only two species were to a small degree capable of autonomous selfing. Protea species varied in their dependence on animal pollinators, including species depending exclusively on birds, insects, or both for seed production. Protea species exhibited continuous variation in the effects of bird-pollinators on reproduction, challenging the accuracy of categorical pollination syndromes for predicting pollinator effects on plant reproduction. The meta-analysis showed that the quality of floral rewards, especially nectar sugar concentration, predicts the dependence of Protea reproduction on bird pollinators, whereas the signalling and accessibility traits tested in this study were not important. Beyond revealing new insights into the pollination biology of Protea species, our findings underscore the usefulness of experimental approaches on multiple plant species to test the effects of pollinator guilds on plant reproduction and contribute to a trait-based understanding of the functional determinants of plant–pollinator interactions. [ABSTRACT FROM AUTHOR]