Determining whether seed production is pollen limited has been an area of intensive empirical study over the last two decades. Yet current evidence does not allow satisfactory assessment of the causes or consequences of pollen limitation. Here, we critically evaluate existing theory and issues concerning pollen limitation. Our main conclusion is that a change in approach is needed to determine whether pollen limitation reflects random fluctuations around a pollen-resource equilibrium, an adaptation to stochastic pollination environments, or a chronic syndrome caused by an environmental perturbation. We formalize and extend D. Haig and M. Westoby's conceptual model, and illustrate its use in guiding research on the evolutionary consequences of pollen limitation, i.e., whether plants evolve or have evolved to ameliorate pollen limitation. This synthesis also reveals that we are only beginning to understand when and how pollen limitation at the plant level translates into effects on plant population dynamics. We highlight the need for both theoretical and empirical approaches to gain a deeper understanding of the importance of life-history characters, Allee effects, and environmental perturbations in population declines mediated by pollen limitation. Lastly, our synthesis identifies a critical need for research on potential effects of pollen limitation at the community and ecosystem levels.

Quantifying the extent to which seed production is limited by the availability of pollen has been an area of intensive empirical study over the past few decades. Whereas theory predicts that pollen augmentation should not increase seed production, numerous empirical studies report significant and strong pollen limitation. Here, we use a variety of approaches to examine the correlates of pollen limitation in an effort to understand its occurrence and importance in plant evolutionary ecology. In particular, we examine the role of recent ecological perturbations in influencing pollen limitation and discuss the relation between pollen limitation and plant traits. We find that the magnitude of pollen limitation observed in natural populations depends on both historical constraints and contemporary ecological factors.

Competition for pollination has served as a model for the integration of ecological and evolutionary perspectives in the study of species interactions. Its study has elucidated both obvious and more subtle mechanisms, and has documented a range of outcomes. However, the potential for this interaction to inform our understanding of both pure and applied aspects of pollination biology has only begun to be realized.

One Sentence Summary: Empirical evidence from grasslands around the world demonstrates a humped-back relationship between plant species richness and biomass at the 1 m 2 plot scale.Abstract: One of the central problems of ecology is the prediction of species diversity. The humped-back model (HBM) suggests that plant diversity is highest at intermediate levels of productivity; at low productivity few species can tolerate the environmental stresses and at high productivity a small number of highly competitive species dominate. A recent study claims to have comprehensively refuted the HBM. Here we show, using the largest, most geographically diverse dataset ever compiled and specifically built for testing this model that if the conditions are met, namely a wide range in biomass at the 1 m 2 plot level and the inclusion of plant litter, the relationship between plant biomass and species richness is hump shaped, supporting the HBM. Our findings shed new light on the prediction of plant diversity in grasslands, which is crucial for supporting management practices for effective conservation of biodiversity. 4Main Text: The relationship between plant diversity and productivity is a topic of intense debate (1-6). The HBM states that plant species richness peaks at intermediate productivity, taking above-ground biomass as a proxy for annual net primary productivity (ANPP) (7-9). This diversity peak is driven by two opposing processes; in unproductive and disturbed ecosystems where there is low plant biomass, species richness is limited by either stress, such as insufficient water and mineral nutrients, or high levels of disturbance-induced removal of biomass, which few species are able to tolerate. In contrast, in the low disturbance and productive conditions that generate high plant biomass it is competitive exclusion by a small number of highly competitive species that is hypothesized to constrain species richness (7-9). Other mechanisms proposed to explain the unimodal relationship between species richness and productivity include disturbance (10), evolutionary history and dispersal limitation (11,12), and density limitation affected by plant size (13).Different case studies have supported or rejected the HBM, and three separate meta-analyses reached different conclusions (14). This inconsistency may indicate a lack of generality of the HBM, or it may reflect a sensitivity to study characteristics including the type(s) of plant communities considered, the taxonomic scope, the length of the gradient sampled, the spatial grain and extent of analyses (14,15), and the particular measure of net primary productivity (16). Although others would argue (6), we maintain that the question remains whether the HBM serves as a useful and general model for grassland ecosystem theory and management. 5 We quantified the form and strength of the richness-productivity relationship using novel data from a globally-coordinated (17), distributed, scale-standardized and consistently designed survey, in which plant richness and biomass were m...

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Invasive species are frequently regarded as superlative competitors that can vegetatively crowd out natives, but little is known about whether invasives can compete for pollination services with native plants. We hypothesized that, when the showy invasive species Lythrum salicaria (purple loosestrife) was present, pollinator visitation and seed set would be reduced in a native congener, L. alatum (winged loosestrife). To test this hypothesis, we constructed mixed and monospecific plots of the two species. Over two years of study, we found that L. salicaria significantly reduced both pollinator visitation and seed set in L. alatum. Furthermore, pollinators moved frequently between the two plant species, which may cause heterospecific pollen transfer. Thus, reductions in both pollen quantity and pollen quality may reduce L. alatum seed set. If similar patterns occur in the field, invasive plants may be an even greater threat to natives than previously thought.

Summary 1.Pollinators visiting large floral displays may probe several flowers in sequence, leading to geitonogamous (among-flower) self-pollination. To investigate the relationship between floral display size and patterns of pollinator movement, we studied foraging by several pollinator species in four replicate arrays of Mimulus ringens (Scrophulariaceae). In each array displays were trimmed to two, four, eight and 16 flowers per plant. 2. Bees preferred large displays, and probed more flowers in sequence on large than on small displays. However, the proportion of available flowers probed decreased with display, resulting in nearly equal floral visitation rates across treatments. 3. Because pollinators probed more flowers in sequence on large displays, plants with numerous flowers should experience more geitonogamous self-pollination than plants with small displays. 4. In all four treatments, pollinators frequently visited only one flower before leaving the plant. As the first flower probed on a plant cannot receive geitonogamous pollen, this potentially reduces selfing rates for those flowers, compared to flowers probed late in a long visitation sequence on a plant. Such differences among flowers in pollination history should increase variation in geitonogamous self-pollination among fruits within plants. 5. The three most abundant pollinator species differed significantly in behaviours that could influence plant mating patterns, including number of flowers probed per plant; interplant movement distances; and grooming. Variation in foraging patterns was also evident among individuals within species. These subtle differences in response should affect the pollination services provided to plants.

Competition for pollination can be an important factor in plant reproduction, but little attention has been given to the effect of the growing number of invasive plant species on pollination of native species. As a first step in understanding this threat, we used hand pollination to investigate the effects of pollen from an invasive species (Lythrum salicaria) on seed set in a sympatric and co-flowering native congener (L. alatum). Dispersal of fluorescent dyes in the field confirms that pollinators (bumble bees and honey bees) transfer pollen between species. To determine the potential effect of such interspecific pollen transfer on seed set of the native, we pollinated 773 flowers on 20 plants with one of three treatments: legitimate conspecific pollen, a mixture of conspecific and foreign pollen, and foreign pollen. The mixed-pollen treatment resulted in 28.8% lower seed set relative to conspecific pollination. Foreign crosses resulted in extremely low seed set. Observations of pollen germination indicate that events at the stigmatic surface contribute to the reduction in seed set for mixed pollination. Our results indicate that the impacts of invasive species may extend beyond vegetative competition to include competition for pollination.