Explore the vast range of titles available.

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.

A generalist pollination system may be characterized through the interaction of a plant species with two or more functional groups of pollinators. The spatiotemporal variation of the most effective pollinator is the factor most frequently advocated to explain the emergence and maintenance of generalist pollination systems. There are few studies merging variation in floral visitor assemblages and the efficacy of pollination by different functional groups. Thus, there are gaps in our knowledge about the variation in time of pollinator efficacy and frequency of generalist species. In this study, we evaluated the pollination efficacy of the floral visitors of Edmundoa lindenii (Bromeliaceae) and their frequency of visits across four reproductive events. We analyzed the frequency of the three groups of floral visitors (large bees, small bees, and hummingbirds) through focal observations in the reproductive events of 2015, 2016, 2017, and 2018. We evaluated the pollination efficacy (fecundity after one visit) through selective exposure treatments and the breeding system by manual pollinations. We tested if the reproductive success after natural pollination varied between the reproductive events and also calculated the pollen limitation index. E. lindenii is a self-incompatible and parthenocarpic species, requiring the action of pollinators for sexual reproduction. Hummingbirds had higher efficacy than large bees and small bees acted only as pollen larcenists. The relative frequency of the groups of floral visitors varied between the reproductive events. Pollen limitation has occurred only in the reproductive event of 2017, when visits by hummingbirds were scarce and reproductive success after natural pollination was the lowest. We conclude that hummingbirds and large bees were the main and the secondary pollinators of E. lindenii , respectively, and that temporal variations in the pollinator assemblages had effects on its reproductive success. Despite their lower pollination efficacy, large bees ensured seed set when hummingbirds failed. Thus, we provide evidence that variable pollination environments may favor generalization, even under differential effectiveness of pollinator groups if secondary pollinators provide reproductive assurance.

1. The best-of-both-worlds hypothesis proposes that delayed autonomous selfing assures reproduction when pollinators are absent, but allows outcrossing when pollinators are present without incurring seed and pollen discounting. However, because self-compatibility is necessary for delayed selfing, discounting can be incurred if pollinators mediate selfing concurrently with outcrossing. The extent that pollinator-mediated selfing erodes the best-of-both-worlds mating strategy is largely unknown. 2. In this study, we assess how delayed selfing and pollinator-mediated selfing contribute to reproductive output under contrasting pollination environments in Bulbine vagans (Asphodelaceae). 3. We manipulated floral phenotype under contrasting weather conditions, plant densities and floral display. We emasculated flowers at early-flowering (allowing only outcrossing) or late-flowering (eliminating delayed selfing) stages and compared reproductive output with intact flowers (outcrossing + pollinator-mediated selfing + delayed selfing). 4. During inclement weather, few or no pollinators visited flowers. Reproductive output was severely limited by cross pollen, and delayed selfing provided reproductive assurance. During fine weather, cross pollen was also limiting, but pollinator-mediated selfing boosted reproductive output before delayed selfing could occur. 5. On isolated plants, flowers received fewer pollinator visits and reproductive output was limited more by cross pollen compared with grouped plants. Despite these differences, reproductive output of both isolated and grouped plants was boosted to similar levels by pollinator-mediated selfing before delayed selfing could occur. 6. One- and three-flowered plants were similarly limited by cross pollen. On three-flowered plants, facilitated and geitonogamous selfing boosted reproductive output before delayed selfing could occur. By contrast, on one-flowered plants geitonogamy was eliminated, and pollinator-mediated selfing was reduced. This provided mating flexibility by retaining opportunities for subsequent outcrossing, although delayed selfing boosted reproductive output. 7. Reproductive output was consistently limited by cross-pollination. Yet, delayed selfing provided reproductive assurance only during inclement weather and on one-flowered plants. Under other conditions, pollinators mediated selfing that could render ovules unavailable for outcrossing as selfing occurred and for outcrossing that might occur subsequently. We conclude that best-of-both-worlds flexibility will often be eroded by pollinator-mediated selfing that inevitability occurs with outcrossing.

Pollination success of animal‐pollinated flowers depends on rate of pollinator visits and on pollen deposition per visit, both of which should vary with the pollen and nectar “neighborhoods” of a plant, i.e., with pollen and nectar availability in nearby plants. One determinant of these neighborhoods is per‐flower production of pollen and nectar, which is likely to respond to environmental influences. In this study, we explored environmental effects on pollen and nectar production and on pollination success in order to follow up a surprising result from a previous study: flowers of Ipomopsis aggregata received less pollen in years of high visitation by their hummingbird pollinators. A new analysis of the earlier data indicated that high bird visitation corresponded to drought years. We hypothesized that drought might contribute to the enigmatic prior result if it decreases both nectar and pollen production: in dry years, low nectar availability could cause hummingbirds to visit flowers at a higher rate, and low pollen availability could cause them to deposit less pollen per visit. A greenhouse experiment demonstrated that drought does reduce both pollen and nectar production by I. aggregata flowers. This result was corroborated across 6 yr of variable precipitation and soil moisture in four unmanipulated field populations. In addition, experimental removal of pollen from flowers reduced the pollen received by nearby flowers. We conclude that there is much to learn about how abiotic and biotic environmental drivers jointly affect pollen and nectar production and availability, and how this contributes to pollen and nectar neighborhoods and thus influences pollination success.

Mast seeding is a widespread and widely studied phenomenon. However, the physiological mechanisms that mediate masting events and link them to weather and plant resources are still debated. Here, we explore how masting is affected by plant resource budgets, fruit maturation success, and hormonal coordination of cues including weather and resources. There is little empirical support for the commonly stated hypothesis that plants store carbohydrates over several years to expend in a high-seed year. Plants can switch carbohydrates away from growth in high-seed years, and seed crops are more probably limited by nitrogen or phosphorus. Resources are clearly involved in the proximate mechanisms driving masting, but resource budget (RB) models cannot create masting in the absence of selection because some underlying selective benefit is required to set the level of a ‘full’ seed crop at greater than the annual resource increment. Economies of scale (EOSs) provide the ultimate factor selecting for masting, but EOSs probably always interact with resources, which modify the relationship between weather cues and reproduction. Thus, RB and EOS models are not alternative explanations for masting – both are required. Experiments manipulating processes that affect mast seeding will help clarify the physiological mechanisms that underlie mast seeding.

Field studies in South Africa showed that floral spur length in the Disa draconis complex (Orchidaceae) varies enormously between populations in the southern mountains (means = 32-38 mm), lowland sandplain (mean = 48 mm), and northern mountains (means = 57-72 mm). We tested the hypothesis that divergence in spur length has resulted from selection exerted through pollinator proboscis length. Short-spurred plants in several southern mountain populations, as well as long-spurred plants in one northern mountain population, were pollinated by a horsefly, Philoliche rostrata (Tabanidae), with a proboscis length that varied from 22 to 35 mm among sites. Longspurred plants on the sandplain were pollinated by the tanglewing fly, Moegistorynchus longirostris (Nemestrinidae), which has a very long proboscis (mean = 57 mm). Selection apparently favors long spurs in sandplain plants, as artificial shortening of spurs resulted in a significant decline in pollen receipt and fruit set, although pollinaria removal was not significantly affected. Fruit set in the study populations was limited by pollen availability, which further suggests that selection on spur length occurs mainly through the female component of reproductive success.

Siring success plays a key role in plant evolution and reproductive ecology, and variation among individuals creates an opportunity for selection to act. Differences in male reproductive success can be caused by processes that occur during two stages, the pollination and post-pollination phases of reproduction. In the pollination phase, heritable variation in floral traits and floral display affect pollinator visitation patterns, which in turn affect variation among plants in the amount of pollen exported and deposited on recipient stigmas. In the post-pollination phase, differences among individuals in pollen grain germination success and pollen tube growth may cause realized paternity to differ from patterns of pollen receipt. The maternal plant can also preferentially provision some developing seeds or fruits to further alter variation in siring success. In this review, we describe studies that advance our understanding of the dynamics of the pollination and post-pollination phases, focusing on how variation in male fitness changes in response to pollen limitation. We then explore the interplay between pollination and post-pollination success, and how these processes respond to ecological factors such as pollination intensity. We also identify pressing questions at the intersection of pollination and paternity and describe novel experimental approaches to elucidate the relative importance of pollination and post-pollination factors in determining male reproductive success. The relative contribution of pollination and post-pollination processes to variation in male reproductive success may not be constant, but rather may vary with pollination intensity. Studies that quantify the effects of pollination and post-pollination phases in concert will be especially valuable as they will enable researchers to more fully understand the ecological conditions influencing male reproductive success.

The diversity and abundance of wild insect pollinators have declined in many agricultural landscapes. Whether such declines reduce crop yields, or are mitigated by managed pollinators such as honey bees, is unclear. We found universally positive associations of fruit set with flower visitation by wild insects in 41 crop systems worldwide. In contrast, fruit set increased significantly with flower visitation by honey bees in only 14% of the systems surveyed. Overall, wild insects pollinated crops more effectively; an increase in wild insect visitation enhanced fruit set by twice as much as an equivalent increase in honey bee visitation. Visitation by wild insects and honey bees promoted fruit set independently, so pollination by managed honey bees supplemented, rather than substituted for, pollination by wild insects. Our results suggest that new practices for integrated management of both honey bees and diverse wild insect assemblages will enhance global crop yields.

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.

In animal-pollinated plants, the opportunity for selection and the strength of pollinatormediated selection are expected to increase with the degree of pollen limitation. However, whether differences in pollen limitation can explain variation in pollinator-mediated and net selection among animal-pollinated species is poorly understood. In the present study, we quantified pollen limitation, variance in relative fitness and pollinator-mediated selection on five traits important for pollinator attraction (flowering start, plant height, flower number, flower size) and pollination efficiency (spur length) in natural populations of 12 orchid species. Pollinator-mediated selection was quantified by subtracting estimates of selection gradients for plants receiving supplemental hand-pollination from estimates obtained for open-pollinated control plants. Mean pollen limitation ranged from zero to 0.96. Opportunity for selection, pollinator-mediated selection and net selection were all positively related to pollen limitation, whereas nonpollinator-mediated selection was not. Opportunity for selection varied five-fold, strength of pollinator-mediated selection varied three-fold and net selection varied 1.5-fold among species. Supplemental hand-pollination reduced both opportunity for selection and selection on floral traits. The results show that the intensity of biotic interactions is an important determinant of the selection regime, and indicate that the potential for pollinator-mediated selection and divergence in floral traits is particularly high in species that are strongly pollen-limited.