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Immobility of flowering plants requires them to engage pollen vectors to outcross, introducing considerable inefficiency in the conversion of pollen production into sired seeds. Whether inefficiencies influence the evolution of the relative resource allocation to female and male functions has been debated for more than 40 years. Whereas early models suggested no effect, negative interspecific relations of mean pollen production and pollen : ovule ratios to the proportion of removed pollen that is exported to stigmas (pollen-transfer efficiency) indicate otherwise. Here, we consider theoretically a key condition that determines whether the efficiencies of processes (first derivative of process output with respect to input) affect the evolutionarily stable sex (ESS) allocation. No effect arises if all individuals experience the same efficiency. By contrast, a decline in process efficiency with increasing allocation (diminishing returns) generally reduces the ESS male allocation for a population. Furthermore, differences in the allocation dependence of efficiencies (and hence the ESS sex allocation) among populations/species create a negative relation of realised efficiency to male allocation among species, like that observed empirically. Diminishing returns arise for various processes that affect siring (e.g. pollen export and local pollen competition to fertilise ovules), which may differ in their relative influence on sex allocation among species., (© 2025 The Author(s). New Phytologist © 2025 New Phytologist Foundation.)

Mating success of flowering plants depends strongly on the efficiencies of pollen removal from flowers and its subsequent dispersal to conspecific stigmas. We characterized the economy of pollen dispersal in flowering plants by analysing pollen fates and their correlates for 228 species. The mean percentage of pollen removed from flowers (removal efficiency) varied almost twofold according to the type of pollen-dispersal unit, from less than 45% for orchids and milkweeds with solid pollinia, to greater than 80% for species with granular monads or sectile (segmented) pollinia. The mean percentage of removed pollen reaching stigmas (pollen transfer efficiency, PTE) varied from 2.4% for species with separate monads to 27.0% for orchids with solid pollinia. These values tended to be higher in plants with single pollinator species and in those with non-grooming pollinators. Nectar production increased removal efficiency, but did not influence PTE. Among types of pollen-dispersal units, the net percentage of produced pollen that was dispersed to stigmas varied negatively with removal efficiency and positively with PTE, indicating the relative importance of the latter for overall pollen economy. These findings confirm the key importance of floral traits, particularly pollen packaging, for pollen dispersal outcomes and highlight the under-appreciated pollination efficiency of non-grooming pollinators.

Premise: The relative per-flower production of ovules and pollen varies broadly with angiosperm mating systems, with outcrossing types commonly producing more pollen grains per ovule than selfing types. The evolutionary causes of this variation are contentious, especially the relevance of pollination risk. Resolution of this debate may have been hampered by its focus on pollen:ovule (P:O) ratios rather than on the evolution of pollen and ovule numbers per se., Methods: Using published mean ovule and pollen counts, we analyzed associations with the proportion of removed pollen that reaches stigmas (pollen-transfer efficiency) and differences between pollinator-dependent and autogamous forms within and among species. Analyses involved Bayesian methods that simultaneously considered variation in pollen and ovule numbers and accounted for phylogenetic relatedness. We also assessed the utility of P:O ratios as mating-system proxies and their association with female outcrossing rates., Results: Median pollen number declined consistently with pollen-transfer efficiency among species, whereas median ovule number did not. Similarly, in both intraspecific and interspecific analyses, pollinator-dependent plants produced more pollen than autogamous plants, whereas ovule production did not differ statistically. Distributions of P:O ratios overlapped extensively for self-incompatible and self-compatible species and for different mating-system classes, and P:O ratios correlated weakly with outcrossing rate., Conclusions: Our findings demonstrate that pollinator dependence and pollination efficiency commonly influence the evolution of pollen number per flower but have more limited effects on ovule number. P:O ratios provide ambiguous, possibly misleading, information about mating systems, especially when compared among clades., (© 2023 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

Reproductive systems of flowering plants are evolutionarily fluid, with mating patterns changing in response to shifts in abiotic conditions, pollination systems, and population characteristics. Changes in mating should be particularly evident in species with sexual polymorphisms that become ecologically destabilized, promoting transitions to alternative reproductive systems. Here, we decompose female mating portfolios (incidence of selfing, outcross mate number, and intermorph mating) in eight populations of Primula oreodoxa, a self-compatible insect-pollinated herb. This species is ancestrally distylous, with populations subdivided into two floral morphs that usually mate with each other (disassortative mating). Stages in the breakdown of polymorphism also occur, including "mixed" populations of distylous and homostylous (self-pollinating) morphs and purely homostylous populations. Population morph ratios vary with elevation in association with differences in pollinator availability, providing an unusual opportunity to investigate changes in mating patterns accompanying transitions in reproductive systems. Unexpectedly, individuals mostly outcrossed randomly, with substantial disassortative mating in at most two distylous populations. As predicted, mixed populations had higher selfing rates than distylous populations, within mixed populations, homostyles selfed almost twice as much as the distylous morphs, and homostylous populations exhibited the highest selfing rates. Populations with homostyles outcrossed with fewer mates and mate number varied negatively with population selfing rates. These differences indicate maintenance of distyly at low elevation, transition to monomorphic selfing at high elevation, and uncertain, possibly variable fates at intermediate elevation. By quantifying the earliest changes in mating that initiate reproductive transitions, our study highlights the key role of mating in promoting evolutionary divergence.

Premise: Fecundity and mating outcomes commonly differ among plant populations occupying contrasting environments. If self-pollination occurs primarily among flowers within plants, contrasting reproductive outcomes among populations must reflect environmental effects on plant-pollinator interactions. Specifically, local conditions could affect features of plant phenotypes that influence pollinator behavior, in turn modifying plant reproductive outcomes., Methods: We compared phenotypes, pollinator abundance and behavior, and female fecundity and mating in two meadow populations and two forest populations of Aconitum kusnezoffii within 3 km of each other. Mating outcomes were assessed using microsatellites., Results: Meadow plants generally produced more, shorter ramets with more, larger flowers, but less nectar per flower than forest plants. These differences likely largely represent phenotypic plasticity. Individual bumble bees visited more flowers on forest plants, likely because the more abundant bees in the meadows depleted nectar availability, as indicated by briefer visits to individual flowers. Despite similar fruit set in both habitats, forest plants set more seeds per fruit. Nevertheless, meadow plants produced more seeds overall, owing to sevenfold greater flower production. Consistent with individual bees visiting fewer flowers on meadow plants, more of their seeds were outcrossed. However, the outcrossed seeds of forest plants included more male mates., Conclusions: Reproductive outcomes can vary among populations of animal-pollinated plants as a result of differences in the availability of effective pollinators and environmental effects on plant phenotypes, and their functional consequences for pollinator behavior that governs pollen dispersal., (© 2022 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

While feeding, foragers can alter their environment. Such alteration constitutes ecological niche construction (ENC) if it enables future benefits for the constructor and conspecific individuals. The environmental modification may also affect non-constructing, bystander species, especially if they share resources with constructor species. If so, ENC could confer the constructor species a competitive advantage by both enhancing its foraging returns and reducing those of bystander species. Expectations - (E1) ENC frequency should vary positively with the recent and current density of the constructor species, and (E2) constructors should use modifications disproportionately. In contrast, bystanders should (E3) experience intensified competition for the affected resource, and (E4) exhibit diverse, possibly mitigating, responses to ENC, depending on opportunity and relative benefits. We investigated these expectations in Argentina for competition for Fuchsia magellanica nectar between an invasive bumble bee Bombus terrestris (terr: putative constructor), which often bites holes at the bases of floral tubes to rob nectar, and native B. dahlbomii (dahl: bystander), which normally accesses Fuchsia nectar through the flower mouth (front visits). Robbing holes constitute ENC, as they persist until the 7-day flowers wilt. The dynamics of the incidence of robbed flowers, abundance of both bees and the number and types of their flower visits (front or robbing) were characterised by alternate-day surveys of plants during 2.5 months. After initially accessing Fuchsia nectar via front visits, terr switched to robbing and its abundance on Fuchsia increased 20-fold within 10 days (E2). Correspondingly, the incidence of robbed flowers varied positively with recent and past terr abundance (E1). In contrast, dahl abundance remained low and varied negatively with the incidence of robbed flowers (E3). When terr ceased visiting Fuchsia, dahl abundance increased sixfold within 10 days (E3), possibly because many dahl previously had avoided competition with terr by feeding on other plant species (E4). While terr was present, dahl on Fuchsia used front visits (tolerance) or used existing robbing holes (adoption: E4). The diverse dahl responses suggest partial compensation for competition with terr. ENC alters competitive asymmetry, favouring constructor species. However, bystander responses can partially offset this advantage, perhaps facilitating coexistence., (© 2021 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

In a recent paper, Renner et al. (Oecologia 195:825-831, 2021) concluded, without supporting statistical evidence, that increased density of managed honey-bee hives between 2019 and 2020 intensified competitive effects of honey bees on non-Apis bee species in the Munich Botanic Garden. Analysis of Renner et al.'s observations revealed that, contrary to their assumption, the change in hive numbers did not statistically alter honey-bee visitation to 29 plant species within or between years. Given this consistency, changes in the proportion of non-Apis bees among visitors of the surveyed plant species between years likely represent their responses to reduced overall availability of floral resources during 2020. Thus, Renner et al.'s observations do not provide convincing evidence that honey bees competitively reduced the abundance of non-Apis bees in the Munich Botanic Garden., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

AbstractSiring success of flowering plants depends on the fates of male gametophytes, which compete for access to stigmas, stylar resources, and ovules. Although rarely considered, pollen may often compete during dispersal, affecting the processes required for export to stigmas: pollen pickup, transport, and deposition. We quantified dispersal interference by tracking bee-mediated dispersal of stained Anacamptis morio (Orchidaceae) pollen from individual donor flowers and inferred the affected dispersal mechanisms on the basis of the fit of a process-based model. During individual trials, all recipient flowers were either emasculated, precluding interference with donor pollen, or intact, adding potentially interfering pollen to the pollinator. The presence of competing pollinaria on bees reduced pickup of additional pollinaria, doubled the overall proportion of lost donor pollen, and reduced total pollen export by 27%. Interference specifically increased loss of donor pollen between successive flower visits and variation in deposition among trials, and it likely also reduced pollen contact with stigmas and pollen deposition when contact occurred. Thus, by altering pollen removal, transport, and deposition, male-male interference during pollen dispersal can significantly-and perhaps commonly-limit plant-siring success.

A recent claim that evening primrose flowers adaptively secrete nectar in response to vibrations from hovering bees lacks supporting evidence. The authors fail to demonstrate that bees can access the concealed nectar and that their visits enhance plant fitness. Reanalysis of the authors' data raises additional concerns about their conclusions., (© 2020 John Wiley & Sons Ltd.)

Interactions between mutualists, competitors, and antagonists have contrasting ecological effects that, sustained over generations, can influence micro- and macroevolution. Dissimilar benefits and costs for these interactions should cause contrasting co-diversification patterns between interacting clades, with prevalent co-speciation by mutualists, association loss by competitors, and host switching by antagonists. We assessed these expectations for a local assemblage of 26 fig species (Moraceae: Ficus), 26 species of mutualistic (pollinating), and 33 species of parasitic (galling) wasps (Chalcidoidea). Using newly acquired gene sequences, we inferred the phylogenies for all three clades. We then compared the three possible pairs of phylogenies to assess phylogenetic congruence and the relative frequencies of co-speciation, association duplication, switching, and loss. The paired phylogenies of pollinators with their mutualists and competitors were significantly congruent, unlike that of figs and their parasites. The distributions of macroevolutionary events largely agreed with expectations for mutualists and antagonists. By contrast, that for competitors involved relatively frequent association switching, as expected, but also unexpectedly frequent co-speciation. The latter result likely reflects the heterogeneous nature of competition among fig wasps. These results illustrate the influence of different interspecific interactions on co-diversification, while also revealing its dependence on specific characteristics of those interactions., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

Ecological interaction and adaptation both depend on phenotypic characteristics. In contrast with the common conception of the 'adult' phenotype, plant bodies develop continuously during their lives. Furthermore, the different units (metamers) that comprise plant bodies are not identical copies, but vary extensively within individuals. These characteristics foster recognition of plant phenotypes as dynamic mosaics. We elaborate this conception based largely on a wide-ranging review of developmental, ecological and evolutionary studies of plant reproduction, and identify its utility in the analysis of plant form, function and diversification. An expanded phenotypic conception is warranted because dynamic mosaic features affect plant performance and evolve. Evidence demonstrates that dynamic mosaic phenotypes enable functional ontogeny, division of labour, resource and mating efficiency. In addition, dynamic mosaic features differ between individuals and experience phenotypic selection. Investigation of the characteristics and roles of dynamic and mosaic features of plant phenotypes benefits from considering within-individual variation as a function-valued trait that can be analysed with functional data methods. Phenotypic dynamics and within-individual variation arise despite an individual's genetic uniformity, and develop largely by heterogeneous gene expression and associated hormonal control. These characteristics can be heritable, so that dynamic mosaic phenotypes can evolve and diversify by natural selection., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

For hermaphroditic angiosperms with multiple flowers, the sex roles can be exclusively combined in bisexual flowers (monocliny), strictly separated among different flowers (monoecy), or arrayed in mixtures of bisexual flowers with female flowers (gynomonoecy) or male flowers (andromonoecy). The hypothesized benefits favoring the evolution of these contrasting hermaphroditic sexual systems are typically examined individually, usually by assessing success through only one sex role. We tested predictions of most hypotheses experimentally with an andromonoecious species, Anticlea occidentalis (Melanthiaceae), based on the performance of intact plants (andromonoecy) and those with emasculated bisexual flowers (functionally monoecious) or emasculated male flowers (functionally monoclinous with sterile peripheral flowers). Andromonoecy in this species enables efficient, size-dependent resource allocation, emphasizing female function in large plants. Emasculation revealed that anthers in male flowers promote female mating quality (outcrossing rate and mate diversity), whereas anthers in bisexual flowers promote male mating quantity (pollen dispersal distance and probability of any siring success). Thus, different hermaphroditic sexual systems likely evolve to capitalize on suites of benefits, rather than just one, and provide compromises between quantitative and qualitative reproductive components. These compromises apparently maximize an individual's combined genetic contributions through female and male functions, rather than separate contributions through each sex role., (© 2018 The Author(s). Evolution © 2018 The Society for the Study of Evolution.)

Globally, agriculture increasingly depends on pollinators to produce many seed and fruit crops. However, what constitutes optimal pollination service for pollinator-dependent crops remains unanswered. We developed a simulation model to identify the optimal pollination service that maximizes fruit quality in crops. The model depicts the pollination (i.e., autonomous self-fertilization, pollen deposition) and post-pollination (i.e., pollen germination, and time from germination to ovule fertilization) processes leading to fruit and seed set and allows for negative flower-pollinator interactions, specifically pistil damage. We parameterized and validated the model based on empirical observations of commercial raspberry in western Argentina. To assess the effects of pollination intensity for fruit production, we conducted simulations over a range of visit number per flower by the two primary managed pollinators worldwide, Apis mellifera and Bombus terrestris. Simulations identified that ~15-35 visits per flower by A. mellifera or ~10-20 visits by B. terrestris provide adequate pollination and maximize raspberry fruit quality (i.e., estimated as the proportion of ovules that develop into drupelets). Visits in excess of these optima reduce simulated fruit quality, and flowers receiving >670 honey bee visits or >470 bumble bee visits would produce fruits of poorer quality than those receiving no bee visits. The simulations generated consistent, unbiased predictions of fruit quality for 12 raspberry fields. This model could be adapted easily to other animal-pollinated crops and used to guide efficient pollinator management in any agro-ecosystem., (© 2018 by the Ecological Society of America.)

Background and Aims: Plant species differ greatly in the three-dimensional arrangements of their flowers (inflorescence architecture). However, the nature of selection responsible for this diversity is poorly understood. Studies that examine among-species variation suggest that inflorescence architecture affects pollinator behaviour, and so should influence plant mating. However, few studies consider the consequences of within-population architectural variation for pollinator behaviour., Methods: We manipulated inflorescence architecture of Delphinium glaucum to contrast bumblebee responses to normal and one-sided (secund) inflorescences., Key Results: The 'dimensionality' of manipulated inflorescences did not affect the number of flowers that bees visited; however, bees moved upward proportionally more on secund inflorescences., Conclusions: This study shows that realistic within-population variation in inflorescence architecture can manipulate pollinator behaviour. These results bear important consequences for plant mating success and the coordinated evolution of inflorescence architecture and floral specialization within inflorescences. These results also question why secund inflorescences are rare, for which we propose four testable explanations., (© The Author 2016. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Why are some traits and trait combinations exceptionally common across the tree of life, whereas others are vanishingly rare? The distribution of trait diversity across a clade at any time depends on the ancestral state of the clade, the rate at which new phenotypes evolve, the differences in speciation and extinction rates across lineages, and whether an equilibrium has been reached. Here we examine the role of transition rates, differential diversification (speciation minus extinction) and non-equilibrium dynamics on the evolutionary history of angiosperms, a clade well known for the abundance of some trait combinations and the rarity of others. Our analysis reveals that three character states (corolla present, bilateral symmetry, reduced stamen number) act synergistically as a key innovation, doubling diversification rates for lineages in which this combination occurs. However, this combination is currently less common than predicted at equilibrium because the individual characters evolve infrequently. Simulations suggest that angiosperms will remain far from the equilibrium frequencies of character states well into the future. Such non-equilibrium dynamics may be common when major innovations evolve rarely, allowing lineages with ancestral forms to persist, and even outnumber those with diversification-enhancing states, for tens of millions of years., (© 2016 The Author(s).)

The fate of male gametophytes after pollen reaches stigmas links pollination to ovule fertilisation, governing subsequent siring success and seed production. Although male gametophyte performance primarily involves cellular processes, an ecological analogy may expose insights into the nature and implications of male gametophyte success. We elaborate this analogy theoretically and present empirical examples that illustrate associated insights. Specifically, we consider pollen loads on stigmas as localised populations subject to density-independent mortality and density-dependent processes as they traverse complex stylar environments. Different combinations of the timing of pollen-tube access to limiting stylar resources (simultaneous or sequential), the tube distribution among resources (repulsed or random) and the timing of density-independent mortality relative to competition (before or after) create signature relations of mean pollen-tube success and its variation among pistils to pollen receipt. Using novel nonlinear regression analyses (two-moment regression), we illustrate contrasting relations for two species, demonstrating that variety in these relations is a feature of reproductive diversity among angiosperms, rather than merely a theoretical curiosity. Thus, the details of male gametophyte ecology should shape sporophyte reproductive success and hence the dynamics and structure of angiosperm populations., (© 2016 John Wiley & Sons Ltd/CNRS.)

Many broadcast spawning corals in multiple reef regions release their gametes with incredible temporal precision just once per year, using the lunar cycle to set the night of spawning. Moonlight, rather than tides or other lunar-regulated processes, is thought to be the proximate factor responsible for linking the night of spawning to the phase of the Moon. We compared patterns of gene expression among colonies of the broadcast spawning coral Acropora millepora at different phases of the lunar cycle, and when they were maintained under one of three experimentally simulated lunar lighting treatments: i) lunar lighting conditions matching those on the reef, or lunar patterns mimicking either ii) constant full Moon conditions, or iii) constant new Moon conditions. Normal lunar illumination was found to shift both the level and timing of clock gene transcription cycles between new and full moons, with the peak hour of expression for a number of genes occurring earlier in the evening under a new Moon when compared to a full Moon. When the normal lunar cycle is replaced with nighttime patterns equivalent to either a full Moon or a new Moon every evening, the normal monthlong changes in the level of expression are destroyed for most genes. In combination, these results indicate that daily changes in moonlight that occur over the lunar cycle are essential for maintaining normal lunar periodicity of clock gene transcription, and this may play a role in regulating spawn timing. These data also show that low levels of light pollution may have an impact on coral biological clocks., (© 2016 Marine Biological Laboratory.)

Premise of Study: Pollen on a stigma represents a local population of male gametophytes vying for access to female gametophytes in the associated ovary. As in most populations, density-independent and density-dependent survival depend on intrinsic characteristics of male gametophytes and environmental (pistil) conditions. These characteristics and conditions could differ among flowers, plants, populations, and species, creating diverse male-gametophyte population dynamics, which can influence seed siring and production., Methods: For nine species, we characterized the relations of both the mean and standard deviation of pollen-tube number at the style base to pollen receipt with nonlinear regression. Models represented asymptotic or peaked relations, providing information about the incidence and magnitude of facilitation and competition, the spatial and temporal characteristics of competition, and the intensity and relative timing of density-independent mortality., Key Results: We infer that pollen tubes of most species competed sequentially, their tips ceasing growth if earlier tubes had depleted stylar space/resources; although two species experienced simultaneous competition. Tube success of three species revealed positive density dependence (facilitation) at low density. For at least four species, density-independent mortality preceded competition. Tube success varied mostly within plants, rather than among plants or conspecific populations. Pollen quality influenced tube success for two of three species; affecting density-independent survival in one and density-dependent performance in the other., Conclusions: The diverse relations of pollen-tube success to pollen receipt evident among just nine species indicate significant contributions of the processes governing pollen germination and tube growth to the reproductive diversity of angiosperms., (© 2016 Botanical Society of America.)

Variation in resource availability can select for traits that reduce the negative impacts of this variability on mean fitness. Such selection may be particularly potent for seed production in flowering plants, as they often experience variation in pollen receipt among individuals and among flowers within individuals. Using analytically tractable models, we examine the optimal allocations for producing ovules, attracting pollen, and maturing seeds in deterministic and stochastic pollen environments. In deterministic environments, the optimal strategy attracts sufficient pollen to fertilize every ovule and mature every zygote into a seed. Stochastic environments select for allocations proportional to the risk of seed production being limited by zygotes or seed maturation. When producing an ovule is cheap and maturing a seed is expensive, among-plant variation selects for attracting more pollen at the expense of producing fewer ovules and having fewer resources for seed maturation. Despite this increased allocation, such populations are likely to be pollen limited. In contrast, within-plant variation generally selects for an overproduction of ovules and, to a lesser extent, pollen attraction. Such populations are likely to be resource limited and exhibit low seed-to-ovule ratios. These results highlight the importance of multiscale variation in the evolution and ecology of resource allocations.

Local adaptation occurs when different environments are dominated by different specialist genotypes, each of which is relatively fit in its local conditions and relatively unfit under other conditions. Analogously, ecological species sorting occurs when different environments are dominated by different competing species, each of which is relatively fit in its local conditions. The simplest theory predicts that spatial, but not temporal, environmental variation selects for local adaptation (or generates species sorting), but this prediction is difficult to test. Although organisms can be reciprocally transplanted among sites, doing so among times seems implausible. Here, we describe a reciprocal transplant experiment testing for local adaptation or species sorting of lake bacteria in response to both temporal and spatial variation in water chemistry. The experiment used a -80°C freezer as a "time machine." Bacterial isolates and water samples were frozen for later use, allowing transplantation of older isolates "forward in time" and newer isolates "backward in time." Surprisingly, local maladaptation predominated over local adaptation in both space and time. Such local maladaptation may indicate that adaptation, or the analogous species sorting process, fails to keep pace with temporal fluctuations in water chemistry. This hypothesis could be tested with more finely resolved temporal data., (© 2014 The Author(s). Evolution © 2014 The Society for the Study of Evolution.)

Angiosperms engage in distributed reproduction, producing sex organs in multiple flowers on one or more inflorescences, including on different physical individuals of clonal plants. We investigated the effects of alternative deployments of artificial flowers for pollinator behavior and simulated pollen dispersal. Plants presented 18 flowers on either one inflorescence (1-I plants) or three inflorescences (3-I plants) spaced either closely or widely. Bees often skipped inflorescences on 3-I plants, visiting an average of 1.5 fewer flowers overall than on 1-I plants. In simulations with all flowers receiving and donating pollen, this behavior caused 7% less geitonogamy for 3-I plants, contradicting a common supposition that clonality increases geitonogamy. Bees generally moved upward within inflorescences and downward between inflorescences. Consequently, in simulations, segregation of pollen receipt to lower flowers and pollen donation to upper flowers reduced self-pollination and enhanced pollen export much more for 1-I plants. Nectar volume per flower had little relevant influence on bee behavior. The observed bee responses and simulated mating results suggest that production of multiple inflorescences and clonality promote pollination quality when flowers simultaneously receive and donate pollen, whereas a single large inflorescence is advantageous when segregation of sex roles among flowers reduces geitonogamy effectively.

Background: Most angiosperms present flowers in inflorescences, which play roles in reproduction, primarily related to pollination, beyond those served by individual flowers alone. An inflorescence's overall reproductive contribution depends primarily on the three-dimensional arrangement of the floral canopy and its dynamics during its flowering period. These features depend in turn on characteristics of the underlying branching structure (scaffold) that supports and supplies water and nutrients to the floral canopy. This scaffold is produced by developmental algorithms that are genetically specified and hormonally mediated. Thus, the extensive inflorescence diversity evident among angiosperms evolves through changes in the developmental programmes that specify scaffold characteristics, which in turn modify canopy features that promote reproductive performance in a particular pollination and mating environment. Nevertheless, developmental and ecological aspects of inflorescences have typically been studied independently, limiting comprehensive understanding of the relations between inflorescence form, reproductive function, and evolution., Scope: This review fosters an integrated perspective on inflorescences by summarizing aspects of their development and pollination function that enable and guide inflorescence evolution and diversification., Conclusions: The architecture of flowering inflorescences comprises three related components: topology (branching patterns, flower number), geometry (phyllotaxis, internode and pedicel lengths, three-dimensional flower arrangement) and phenology (flower opening rate and longevity, dichogamy). Genetic and developmental evidence reveals that these components are largely subject to quantitative control. Consequently, inflorescence evolution proceeds along a multidimensional continuum. Nevertheless, some combinations of topology, geometry and phenology are represented more commonly than others, because they serve reproductive function particularly effectively. For wind-pollinated species, these combinations often represent compromise solutions to the conflicting physical influences on pollen removal, transport and deposition. For animal-pollinated species, dominant selective influences include the conflicting benefits of large displays for attracting pollinators and of small displays for limiting among-flower self-pollination. The variety of architectural components that comprise inflorescences enable diverse resolutions of these conflicts.

Background and Aims: The Orchidaceae have a history of recurring convergent evolution in floral function as nectar production has evolved repeatedly from an ancestral nectarless state. However, orchids exhibit considerable diversity in nectary type, position and morphology, indicating that this convergence arose from alternative adaptive solutions. Using the genus Disa, this study asks whether repeated evolution of floral nectaries involved recapitulation of the same nectary type or diversifying innovation. Epidermis morphology of closely related nectar-producing and nectarless species is also compared in order to identify histological changes that accompanied the gain or loss of nectar production., Methods: The micromorphology of nectaries and positionally equivalent tissues in nectarless species was examined with light and scanning electron microscopy. This information was subjected to phylogenetic analyses to reconstruct nectary evolution and compare characteristics of nectar-producing and nectarless species., Key Results: Two nectary types evolved in Disa. Nectar exudation by modified stomata in floral spurs evolved twice, whereas exudation by a secretory epidermis evolved six times in different perianth segments. The spur epidermis of nectarless species exhibited considerable micromorphological variation, including strongly textured surfaces and non-secreting stomata in some species. Epidermis morphology of nectar-producing species did not differ consistently from that of rewardless species at the magnifications used in this study, suggesting that transitions from rewardlessness to nectar production are not necessarily accompanied by visible morphological changes but only require sub-cellular modification., Conclusions: Independent nectary evolution in Disa involved both repeated recapitulation of secretory epidermis, which is present in the sister genus Brownleea, and innovation of stomatal nectaries. These contrasting nectary types and positional diversity within types imply weak genetic, developmental or physiological constraints in ancestral, nectarless Disa. Such functional convergence generated by morphologically diverse solutions probably also underlies the extensive diversity of nectary types and positions in the Orchidaceae.

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.

The limits on annual seed production have long been characterized as restriction by either pollination success or resource provision to seed development. This expected dichotomy between pollen and resource limitation is based on the assumption that reproductive resources are fixed, which is reasonable for semelparous species. In contrast, iteroparity can ease the constraints on reproductive output per breeding season, if resources can be either mobilized from past storage or borrowed against future performance. For perennial plants, these options allow enhanced reproductive investment in response to unusually good pollination, so that annual seed production may not be pollen or resource limited. We assessed demand-governed reproductive investment by manipulating both resource supply capacity (partial defoliation) and resource demand (pollination quality: fully self-pollination, fully cross-pollination, or combinations of partial self- and cross-pollination within the inflorescence) for a forest herb, Stenanthium occidentale, which is subject to strong pre-dispersal inbreeding depression. Insensitivity to partial defoliation indicated that reproductive output was not source regulated. Instead, demand by developing seeds governs resource distribution, as demonstrated by elevated photosynthate translocation to fruits on fully cross-pollinated plants and the ability of completely defoliated plants to produce seeds. Such contingent resource allocation eliminates a simple dichotomy between pollen receipt and resource availability as limits on annual seed production. Instead, such flexible reproductive investment allows iteroparous perennials to participate maximally in current reproduction (as determined by ovule production) following superior pollination, or to conserve resources for future reproduction following poor pollination.

Background and Aims: Large floral displays have opposing consequences for animal-pollinated angiosperms: they attract more pollinators but also enable elevated among-flower self-pollination (geitonogamy). The presence of sterile flowers as pollinator signals may enhance attraction while allowing displays of fewer open fertile flowers, limiting geitonogamy. The simultaneous contributions of fertile and non-fertile display components to pollinator attraction and reproductive output remain undetermined., Methods: The simultaneous effects of the presence of sterile flowers and fertile-flower display size in two populations of Leopoldia comosa were experimentally assessed. Pollinator behaviour, pollen removal and deposition, and fruit and seed production were compared between intact plants and plants with sterile flowers removed., Key Results: The presence of sterile flowers almost tripled pollinator attraction, supplementing the positive effect of the number of fertile flowers on the number of bees approaching inflorescences. Although attracted bees visited more flowers on larger inflorescences, the number visited did not additionally depend on the presence of sterile flowers. The presence of sterile flowers improved all aspects of plant performance, the magnitude of plant benefit being context dependent. During weather favourable to pollinators, the presence of sterile flowers increased pollen deposition on stigmas of young flowers, but this difference was not evident in older flowers, probably because of autonomous self-pollination in poorly visited flowers. Total pollen receipt per stigma decreased with increasing fertile display size. In the population with more pollinators, the presence of sterile flowers increased fruit number but not seed set or mass, whereas in the other population sterile flowers enhanced seeds per fruit, but not fruit production. These contrasts are consistent with dissimilar cross-pollination and autonomous self-pollination, coupled with the strong predispersal inbreeding depression exhibited by L. comosa populations., Conclusions: Sterile flowers enrich pollination quality by promoting pollen export and import, while limiting the mating costs of geitonogamy associated with large fertile displays.

Inbreeding depression can reduce the performance of offspring produced by mating between relatives, with consequences for population dynamics and sexual-system evolution. In flowering plants, inbreeding depression commonly acts most intensely during seed development. This predispersal component is typically estimated by comparing seed production following exclusive self- and cross-pollination, but such estimates are unbiased only if seed production is limited by ovule availability, rather than by pollen receipt or seed-development resources. To overcome this problem, we propose experimental and statistical methods based on a model of ovule fertilization and seed development that accounts for differential fertilization by self- and cross-pollen, limited ovule viability or receptivity, differential survival of self- and cross-zygotes and limited resource availability. Simulations illustrate that the proposed methods eliminate bias in estimated predispersal inbreeding depression caused by pollen limitation and can improve estimates under resource limitation. Application of these methods to two orchid species further demonstrates their utility in identifying and estimating diverse influences on reproductive performance under typical conditions. Although our theoretical results raise questions about the reported intensity of predispersal inbreeding depression, our proposed methods guard against bias while also providing insight into plant reproduction., (© 2011 The Author(s). Evolution© 2011 The Society for the Study of Evolution.)

Most rewardless orchids engage in generalized food-deception, exhibiting floral traits typical of rewarding species and exploiting the instinctive foraging of pollinators. Generalized food-deceptive (GFD) orchids compete poorly with rewarding species for pollinator services, which may be overcome by flowering early in the growing season when relatively more pollinators are naive and fewer competing plant species are flowering, and/or flowering for extended periods to enhance the chance of pollinator visits. We tested these hypotheses by manipulating flowering time and duration in a natural population of Calypso bulbosa and quantifying pollinator visitation based on pollen removal. Both early and long flowering increased bumble-bee visitation compared with late and brief flowering, respectively. To identify the cause of reduced visitation during late flowering, we tested whether negative experience with C. bulbosa (avoidance learning) and positive experience with a rewarding species, Arctostaphylos uva-ursi, (associative learning) by captive bumble-bees could reduce C. bulbosa's competitiveness. Avoidance learning explained the higher visitation of early- compared with late-flowering C. bulbosa. The resulting pollinator-mediated selection for early flowering may commonly affect GFD orchids, explaining their tendency to flower earlier than rewarding orchids. For dissimilar deceptive and rewarding sympatric species, associative learning may additionally favour early flowering by GFD species.

Background and Aims: Pollen-collecting bees are among the most important pollinators globally, but are also the most common pollen thieves and can significantly reduce plant reproduction. The pollination efficiency of pollen collectors depends on the frequency of their visits to female(-phase) flowers, contact with stigmas and deposition of pollen of sufficient quantity and quality to fertilize ovules. Here we investigate the relative importance of these components, and the hypothesis that floral and inflorescence characteristics mediate the pollination role of pollen collection by bees., Methods: For ten Aloe species that differ extensively in floral and inflorescence traits, we experimentally excluded potential bird pollinators to quantify the contributions of insect visitors to pollen removal, pollen deposition and seed production. We measured corolla width and depth to determine nectar accessibility, and the phenology of anther dehiscence and stigma receptivity to quantify herkogamy and dichogamy. Further, we compiled all published bird-exclusion studies of aloes, and compared insect pollination success with floral morphology., Key Results: Species varied from exclusively insect pollinated, to exclusively bird pollinated but subject to extensive pollen theft by insects. Nectar inaccessibility and strong dichogamy inhibited pollination by pollen-collecting bees by discouraging visits to female-phase (i.e. pollenless) flowers. For species with large inflorescences of pollen-rich flowers, pollen collectors successfully deposited pollen, but of such low quality (probably self-pollen) that they made almost no contribution to seed set. Indeed, considering all published bird-exclusion studies (17 species in total), insect pollination efficiency varied significantly with floral shape., Conclusions: Species-specific floral and inflorescence characteristics, especially nectar accessibility and dichogamy, control the efficiency of pollen-collecting bees as pollinators of aloes.

Background: The production of flowers, fruits and seeds demands considerable energy and nutrients, which can limit the allocation of these resources to other plant functions and, thereby, influence survival and future reproduction. The magnitude of the physiological costs of reproduction depends on both the factors limiting seed production (pollen, ovules or resources) and the capacity of plants to compensate for high resource demand., Methods: To assess the magnitude and consequences of reproductive costs, we used shading and defoliation to reduce photosynthate production by fully pollinated plants of a perennial legume, Oxytropis sericea (Fabaceae), and examined the resulting impact on photosynthate allocation, and nectar, fruit and seed production., Key Results: Although these leaf manipulations reduced photosynthesis and nectar production, they did not alter photosynthate allocation, as revealed by (13)C tracing, or fruit or seed production. That photosynthate allocation to reproductive organs increased >190 % and taproot mass declined by 29 % between flowering and fruiting indicates that reproduction was physiologically costly., Conclusions: The insensitivity of fruit and seed production to leaf manipulation is consistent with either compensatory mobilization of stored resources or ovule limitation. Seed production differed considerably between the two years of the study in association with contrasting precipitation prior to flowering, perhaps reflecting contrasting limits on reproductive performance.

To communicate with animals, plants use signals that are distinct from their surroundings. Animals generally learn to use these signals through associative conditioning; however, signals are most effective when they elicit innate behavioural responses. Many plant species have flowers specialized for pollination by ground-dwelling mammals, but the signals used to attract these pollinators have not been elucidated. Here, we demonstrate the chemical basis for attraction of mammal pollinators to flowers of the dioecious parasitic plant Cytinus visseri (Cytinaceae). Two aliphatic ketones dominate the scent of this species; 3-hexanone, which elicits strong innate attraction in rodents, and 1-hexen-3-one, which repels them in isolation, but not in combination with 3-hexanone. The aliphatic ketone-dominated scent of C. visseri contrasts with those of insect-pollinated plants, which are typically dominated by terpenoids, aromatic or non-ketone aliphatic compounds. 3-hexanone is also known from some bat-pollinated species, suggesting independent evolution of plant signals in derived, highly specialized mammal-pollination systems.

Human welfare depends on the amount and stability of agricultural production, as determined by crop yield and cultivated area. Yield increases asymptotically with the resources provided by farmers' inputs and environmentally sensitive ecosystem services. Declining yield growth with increased inputs prompts conversion of more land to cultivation, but at the risk of eroding ecosystem services. To explore the interdependence of agricultural production and its stability on ecosystem services, we present and test a general graphical model, based on Jensen's inequality, of yield-resource relations and consider implications for land conversion. For the case of animal pollination as a resource influencing crop yield, this model predicts that incomplete and variable pollen delivery reduces yield mean and stability (inverse of variability) more for crops with greater dependence on pollinators. Data collected by the Food and Agriculture Organization of the United Nations during 1961-2008 support these predictions. Specifically, crops with greater pollinator dependence had lower mean and stability in relative yield and yield growth, despite global yield increases for most crops. Lower yield growth was compensated by increased land cultivation to enhance production of pollinator-dependent crops. Area stability also decreased with pollinator dependence, as it correlated positively with yield stability among crops. These results reveal that pollen limitation hinders yield growth of pollinator-dependent crops, decreasing temporal stability of global agricultural production, while promoting compensatory land conversion to agriculture. Although we examined crop pollination, our model applies to other ecosystem services for which the benefits to human welfare decelerate as the maximum is approached.

Pollen is unique among floral rewards in functioning as both a carrier of gametes and an attractant and nutritious resource for floral visitors. Animals that collect pollen without pollinating (pollen thieves) could reduce siring success of thieved plants and cause pollen limitation of seed set at the population level; however, such impacts on plant reproduction have not been demonstrated experimentally. To test these effects we added hives of native honey bees (Apis mellifera scutellata) to populations of a primarily bird-pollinated plant, Aloe maculata, in eastern South Africa. In field and aviary trials, bee addition increased pollen removal from anthers but decreased pollen deposition on stigmas, and so reduced both male and female pollination components. Further, total seed production decreased with hive addition in the aviary experiment and in three of four field populations, indicating that population-level pollen theft can also compromise reproductive success. In the field, naturally occurring allodapine bees also seemed to act as pollen thieves, outweighing the effect of honey bee hive addition at one of the four aloe populations. Our results highlight the importance of social bees as pollen thieves, even of plants that have evolved in their presence, and the role of dichogamy in promoting pollen theft. Given the commonness of both social bees and dichogamy, pollen theft is likely a much more common influence on floral ecology and evolution than suggested by the sparse literature.

Pollen limitation (PL) of seed production creates unique conditions for reproductive adaptation by angiosperms, in part because, unlike under ovule or resource limitation, floral interactions with pollen vectors can contribute to variation in female success. Although the ecological and conservation consequences of PL have received considerable attention in recent times, its evolutionary implications are poorly appreciated. To identify general influences of PL on reproductive adaptation compared with those under other seed-production limits and their implications for evolution in altered environments, we derive a model that incorporates pollination and post-pollination aspects of PL. Because PL always favours increased ovule fertilization, even when population dynamics are not seed limited, it should pervasively influence selection on reproductive traits. Significantly, under PL the intensity of inbreeding does not determine whether outcrossing or autonomous selfing can evolve, although it can affect which response is most likely. Because the causes of PL are multifaceted in both natural and anthropogenically altered environments, the possible outcrossing solutions are diverse and context dependent, which may contribute to the extensive variety of angiosperm reproductive characteristics. Finally, the increased adaptive options available under PL may be responsible for positive global associations between it and angiosperm diversity.

Declines in the stocks of domesticated honey bees in some countries have been attributed to disease, which is at odds with an increasing global trend in the total number of hives. Based on data on annual growth rates in hive numbers and honey production for 87 countries, we tested the hypothesis that geographic heterogeneity in the growth of the domesticated honey bee population can be attributed to disease. In contrast to predictions of this hypothesis, changes in honey production varied in proportion to changes in hive number. Also, growth in honey production was not more spatially heterogeneous than growth in hive numbers, as expected under a scenario of contagious pests. We argue that although disease aggravates production costs, it has less effect on changes in national hive numbers than labor costs, so that geographic variation in the growth of the global honey bee stock reflects the global division of human labor that is a hallmark of economic globalization, rather than persistent and pervasive biological causes.

Pollen dispersal by animals varies extensively because of differences in pollinator visitation rates among plants, dissimilar pollination by the various pollinators that visit individual plants, and stochastic variation in deposition as an individual pollinator disperses a plant's pollen to subsequently visited recipient flowers. Such variation reduces expected female and male success if seed production decelerates with increasing pollen receipt, because less than average receipt diminishes mean seed production more than copious pollination increases it (Jensen's inequality). We report empirical studies of the nature and magnitude of pollen dispersal variance, which provide the basis for a numerical model of the consequences of dispersal for expected seed production. Model fitting revealed that dispersal of Brassica napus pollen by bumblebees and especially butterflies exhibited much more variation than is expected of a binomial process and was best modeled as a beta-binomial process with a constant mean. Overdispersion arose primarily during pollen dispersal by individual insects, since differences between individuals of the same pollinator type were limited. Our model revealed variance limitation as a previously unrecognized, substantial, and ubiquitous component of pollen limitation of seed production. Variance limitation should select for floral traits that increase pollinator visitation, reduce dispersal variance, or reduce the postpollination nonlinearities that cause Jensen's inequality.

Although not 'a professed botanist', Charles Darwin made seminal contributions to understanding of floral and inflorescence function while seeking evidence of adaptation by natural selection. This review considers the legacy of Darwin's ideas from three perspectives. First, we examine the process of floral and inflorescence adaptation by surveying studies of phenotypic selection, heritability and selection responses. Despite widespread phenotypic and genetic capacity for natural selection, only one-third of estimates indicate phenotypic selection. Second, we evaluate experimental studies of floral and inflorescence function and find that they usually demonstrate that reproductive traits represent adaptations. Finally, we consider the role of adaptation in floral diversification. Despite different diversification modes (coevolution, divergent use of the same pollen vector, pollinator shifts), evidence of pollination ecotypes and phylogenetic patterns suggests that adaptation commonly contributes to floral diversity. Thus, this review reveals a contrast between the inconsistent occurrence of phenotypic selection and convincing experimental and comparative evidence that floral traits are adaptations. Rather than rejecting Darwin's hypotheses about floral evolution, this contrast suggests that the tempo of creative selection varies, with strong, consistent selection during episodes of diversification, but relatively weak and inconsistent selection during longer, 'normal' periods of relative phenotypic stasis.

The prospect that a global pollination crisis currently threatens agricultural productivity has drawn intense recent interest among scientists, politicians, and the general public. To date, evidence for a global crisis has been drawn from regional or local declines in pollinators themselves or insufficient pollination for particular crops. In contrast, our analysis of Food and Agriculture Organization (FAO) data reveals that the global population of managed honey-bee hives has increased approximately 45% during the last half century and suggests that economic globalization, rather than biological factors, drives both the dynamics of the global managed honey-bee population and increasing demands for agricultural pollination services. Nevertheless, available data also reveal a much more rapid (>300%) increase in the fraction of agriculture that depends on animal pollination during the last half century, which may be stressing global pollination capacity. Although the primary cause of the accelerating increase of the pollinator dependence of commercial agriculture seems to be economic and political and not biological, the rapid expansion of cultivation of many pollinator-dependent crops has the potential to trigger future pollination problems for both these crops and native species in neighboring areas. Such environmental costs merit consideration during the development of agriculture and conservation policies.

Many of the diverse animals that consume floral rewards act as efficient pollinators; however, others 'steal' rewards without 'paying' for them by pollinating. In contrast to the extensive studies of the ecological and evolutionary consequences of nectar theft, pollen theft and its implications remain largely neglected, even though it affects plant reproduction more directly. Here we review existing studies of pollen theft and find that: (1) most pollen thieves pollinate other plant species, suggesting that theft generally arises from a mismatch between the flower and thief that precludes pollen deposition, (2) bees are the most commonly documented pollen thieves, and (3) the floral traits that typically facilitate pollen theft involve either spatial or temporal separation of sex function within flowers (herkogamy and dichogamy, respectively). Given that herkogamy and dichogamy occur commonly and that bees are globally the most important floral visitors, pollen theft is likely a greatly under-appreciated component of floral ecology and influence on floral evolution. We identify the mechanisms by which pollen theft can affect plant fitness, and review the evidence for theft-induced ecological effects, including pollen limitation. We then explore the consequences of pollen theft for the evolution of floral traits and sexual systems, and conclude by identifying key directions for future research.

Hermaphroditism allows considerable scope for contributing genes to subsequent generations through various mixtures of selfed and outcrossed offspring. The fitness consequences of different family compositions determine the evolutionarily stable mating strategy and depend on the interplay of genetic features, the nature of mating, and factors that govern offspring development. This theoretical article considers the relative contributions of these influences and their interacting effects on mating-system evolution, given a fixed genetic load within a population. Strong inbreeding depression after offspring gain independence selects for exclusive outcrossing, regardless of the intensity of predispersal inbreeding depression, unless insufficient mating limits offspring production. The extent to which selfing evolves under weak postdispersal inbreeding depression depends on predispersal inbreeding depression and the opportunity for resource limitation of offspring production. Mixed selfing and outcrossing is an evolutionarily stable strategy (ESS) if selfed zygotes survive poorly, but selfed offspring survive well, and maternal individuals produce enough "extra" eggs that deaths of unviable outcrossed embryos do not impact offspring production (reproductive compensation). Mixed mating can also be an ESS, despite weak lifetime inbreeding depression, if self-mating reduces the number of male gametes available for outcrossing (male-gamete discounting). Reproductive compensation and male-gamete discounting act largely independently on mating-system evolution. ESS mating systems always involve either complete fertilization or fertilization of enough eggs to induce resource competition among embryos, so although reproductive assurance is adaptive with insufficient mating, it is never an ESS. Our results illustrate the theoretical importance of different constraints on offspring production (availability of male gametes, egg production, and maternal resources) for both the course and outcome of mating-system evolution, whereas unequal competition between selfed and outcrossed embryos has limited effect. These results also underscore the significance of heterogeneity in the nature and intensity of inbreeding depression during the life cycle for the evolution of hermaphrodite mating systems.

To understand the constraints on biological diversity, we analyzed how selection and development interact to control the evolution of inflorescences, the branching structures that bear flowers. We show that a single developmental model accounts for the restricted range of inflorescence types observed in nature and that this model is supported by molecular genetic studies. The model predicts associations between inflorescence architecture, climate, and life history, which we validated empirically. Paths, or evolutionary wormholes, link different architectures in a multidimensional fitness space, but the rate of evolution along these paths is constrained by genetic and environmental factors, which explains why some evolutionary transitions are rare between closely related plant taxa.

Unlike pollen and seed size, the extent and causes of variation in ovule size remain unexplored. Based on 45 angiosperm species, we assessed whether intra- and interspecific variation in ovule size is consistent with cost minimization during ovule production or allows maternal plants to dominate conflict with their seeds concerning resource investment. Despite considerable intraspecific variation in ovule volume (mean CV = 0.356), ovule production by few species was subject to a size-number trade-off. Among the sampled species, ovule volume varied two orders of magnitude, whereas seed volume varied four orders of magnitude. Ovule volume varied positively among species with flower mass and negatively with ovule number. Tenuinucellate ovules were generally larger that crassinucellate ovules, and species with apical placentation (which mostly have uniovulate ovaries) had smaller ovules than those with other placentation types. Seed volume varied positively among species with fruit mass and seed development time, but negatively with seed number. Seeds grew a median 93-fold larger than the ovules from which they originated. Our results provide equivocal evidence that selection minimizes ovule size to allow efficient resource allocation after fertilization, but stronger evidence that ovule size affords maternal plants an advantage in parent-offspring conflict.

Pollination commonly limits seed production, as addition of pollen to stigmas often increases fecundity. This response is usually interpreted as evidence that plants' stigmas receive too few pollen grains to maximize ovule fertilization (quantity limitation); however, many genetic studies demonstrate that poor-quality pollen can also reduce seed production (quality limitation). We explore both aspects of pollen limitation theoretically with a dose-response model that incorporates a saturating negative-exponential relation of seed production to pollen receipt. This relation depends on aspects of ovule production, pollen import, pollen-pistil interactions and seed development, all of which can contribute to pollen limitation. Our model reveals that quantity limitation is restricted to the lowest range of pollen receipt, for which siring success per pollen grain is high, whereas quality limitation acts throughout the range of pollen receipt if plants do not import the highest-quality pollen. In addition to pollinator availability and efficiency, quantity limitation is governed by all post-pollination aspects of seed production. In contrast, quality limitation depends on the difference in survival of embryos sired by naturally delivered pollen vs. by pollen of maximal quality. We briefly illustrate the distinction between these two components of pollen limitation with results from the mistletoe Tristerix corymbosus. Our model also shows that the standard pollen-supplementation technique neither estimates the total intensity of pollen limitation nor distinguishes between its quantity and quality components. As an alternative, we propose a methodological protocol that requires both measurement of seed production following excess pollination with only outcross pollen and quantification of the dose-response relation of seed output to pollen receipt. This method estimates both the total extent of pollen limitation and its two components. Finally, we consider the influences on quantity and quality limitation, which reveals that quantity limitation probably occurs much less often than has been inferred from pollen-supplementation experiments. These interpretations suggest that an expanded perspective that recognizes the fecundity consequences of pollination with poor-quality pollen would promote ecological understanding of pollen limitation.

Angiosperms display flowers in many three-dimensional arrangements, but the functional significance of this diversity is largely unknown. We examined influences of inflorescence architecture on pollination and mating by quantifying the responses of bumblebees to three architectures and then using these observations as the basis of a model that simulated pollen dispersal. On artificial panicles, racemes, and umbels, each with 12 identical flowers, bees visited one more flower, on average, on umbels than on panicles (with racemes being intermediate). In contrast to this weak response, the consistency of foraging paths among flowers differed strongly among architectures (raceme > panicle > umbel). The simulation model revealed limited differences in self-pollination and pollen export among the three inflorescence designs when all flowers presented and received pollen, because mating differences depended on only the number of flowers visited. In contrast, in simulations of inflorescences on which pollen receipt and presentation were segregated so as to minimize interference among flowers, the consistency of movement paths governed mating. In this case, racemes self-pollinated much less than umbels (with panicles being intermediate), and racemes exported much more pollen than umbels and panicles. These effects have diverse consequences for the evolution of inflorescence architecture, flower design, and sexual segregation.

Plants need not participate passively in their own mating, despite their immobility and reliance on pollen vectors. Instead, plants may respond to their recent pollination experience by adjusting the number of flowers that they display simultaneously. Such responsiveness could arise from the dependence of floral display size on the longevity of individual flowers, which varies with pollination rate in many plant species. By hand-pollinating some inflorescences, but not others, we demonstrate plasticity in display size of the orchid Satyrium longicauda. Pollination induced flower wilting, but did not affect the opening of new flowers, so that within a few days pollinated inflorescences displayed fewer flowers than unpollinated inflorescences. During subsequent exposure to intensive natural pollination, pollen removal and receipt increased proportionally with increasing display size, whereas pollen-removal failure and self-pollination accelerated. Such benefit-cost relations allow plants that adjust display size in response to the prevailing pollination rate to increase their attractiveness when pollinators are rare (large displays), or to limit mating costs when pollinators are abundant (small displays). Seen from this perspective, pollination-induced flower wilting serves the entire plant by allowing it to display the number of flowers that is appropriate for the current pollination environment.

The aerodynamics of wind pollination selects for an intimate relation between form and function in anemophilous plants. Inflorescence architecture and floral morphology vary extensively within the Poaceae, but the functional implication of this variation remains largely unknown. Here we quantify associations between floret, culm, and inflorescence characteristics for 25 grass species in Kananaskis, Alberta, Canada, and consider whether different architectures may implement unique mechanisms to aid pollination. The species cluster into four categories defined by all combinations of floret size (small vs. large) and inflorescence architecture (compact vs. diffuse). Species differed significantly for all 18 traits that we measured, with 12 traits differing only between floret-size classes, three differing only between inflorescence types, and three differing among both (independently or by an interaction). Based on these morphological associations, we discuss the aerodynamic and functional consequences of each category for wind pollination. The independence of inflorescence and floral traits has probably allowed exploration of all possible combinations of inflorescence architecture and floret size during the evolution of the Poaceae.

Background and Aims: Distyly has been hypothesized to promote cross-pollination by reducing intrafloral and geitonogamous self-pollination, and enhancing intermorph pollination. Distylous plants typically display both reciprocal herkogamy and a heteromorphic incompatibility system, which allows mating only between morphs. Distyly and its pollination consequences were examined in two Pentanisia species with long-tubed flowers which are pollinated almost exclusively by butterflies., Methods: Anther and stigma heights were measured to quantify reciprocal herkogamy. The type of incompatibility system was determined by observing pollen tubes and seed production following controlled hand pollination. Pollen loads on pollinators and stigmas were also examined to assess the efficiency of intermorph pollen flow., Key Results: Pentanisia prunelloides and P. angustifolia exhibit reciprocal herkogamy and a host of ancillary dimorphisms, including pollen colour, exine sculpturing, stigmatic papilla shape and floral-tube pubescence. Controlled hand-pollinations revealed the presence of a strong heteromorphic incompatibility system in both species. The site of incompatibility differed between the morphs; intramorph pollen tubes were blocked in the style of the short-styled morph and on the stigmatic surface of the long-styled morph. Butterflies carried pollen from the short- and long-styled morphs primarily on their head and proboscis, respectively. Natural pollination resulted in a higher proportion of pollen transfer from long- to short-styled plants than vice versa. Nevertheless, fruit set did not differ between morphs., Conclusions: Both Pentanisia species are fully distylous. Reciprocal herkogamy results in pollen from the two morphs being carried on different locations on pollinators' bodies, which in turn promotes intermorph pollination. Intramorph pollination does not result in fertilization, because of an effective heteromorphic incompatibility system.

Narcissus, the daffodil genus, exhibits an unusual diversity of sexual systems, with populations that are monomorphic, dimorphic or trimorphic for style length. Associated with this variation are striking differences among species in floral morphology and pollination biology. This diversity provides an opportunity to investigate the evolution of mating polymorphisms, and to determine how floral morphology promotes transitions among sexual systems. Because of the absence of heteromorphic incompatibility in Narcissus, floral morphology plays a key role in governing patterns of outcrossed mating. Phylogenetic evidence indicates that stylar monomorphism is ancestral in the genus, with multiple origins of stylar polymorphism, including independent origins of stigma-height dimorphism, distyly and tristyly. Sexual polymorphisms have evolved only in lineages with narrow floral tubes that are pollinated by Lepidoptera and/or long-tongued bees. Populations of polymorphic Narcissus species are typically dominated by the long-styled morph and display imperfect reciprocity in the positions of sexual organs. These features are consequences of the unusual association between stylar polymorphism and a self-incompatibility system that permits intramorph mating.

In sexually polymorphic species, reproductive morphology governs mating patterns and the character of negative frequency-dependent selection. If local environmental conditions cause sexual morphs to differ between populations, then frequency-dependent selection should create corresponding geographic variation in morph frequencies. We investigate this relation with a model of morph-ratio evolution and analysis of geographic variation in the heterostylous plant Narcissus triandrus. Unlike other tristylous species, N. triandrus possesses both imperfect reciprocity among morphs in sex-organ position and a self-incompatibility system that permits outcrossing within and between morphs. We sampled 137 populations throughout the Iberian Peninsula for floral-morph ratios, and measured floral morphology in 31 populations. Morph ratios exhibited three atypical features: (1) predominance of the long-styled (L) morph; (2) absence of the mid-styled (M) morph from 17.5% of populations; and (3) a negative relation between the frequencies of the L and M morphs among populations. Morph ratios varied geographically, with decreasing frequency of the M morph from the southeast to the northwest of the species' range. Much of this variation accompanied allometric change in the positions of sex organs, especially the mid-level organs, with the M morph declining in frequency and ultimately being lost in large-flowered populations. Using multivariate multiple regression, we demonstrate that variation in floral morphology among populations predicts this geographic variation in morph frequencies. Our theoretical analysis illustrates that patterns of pollen transfer governed by imperfect sex-organ reciprocity can select for unequal equilibrium morph ratios like those observed for N. triandrus. We interpret the L-biased morph ratios and the unusual morphology of N. triandrus as a consequence of its atypical intramorph compatibility system.