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Premise of research. Floral longevity, the amount of time from anthesis to floral senescence, is a key floral trait that determines the opportunity for pollinator visitation. However, the causes and consequences of variation in longevity have primarily been studied among species, which may not reflect patterns of variation within populations. Methodology. To determine the causes and consequences of within-population variation in floral longevity, we exposed Mimulus guttatus to either an ambient- or reduced-pollination treatment. We measured floral longevity to test whether reduced pollinator access (which simulates pollinator decline) causes a plastic extension in floral longevity. We also measured two traits—flower size and anther-stigma distance—that could cause floral longevity to vary and two traits—daily display size and flowering duration—that could vary in response to longevity. Pivotal results. Floral longevity was not affected by experimentally reducing pollinator access or by anther-stigma distance. However, longevity was reduced in M. guttatus plants with larger flowers. Variation in floral longevity did not affect the flowering duration of M. guttatus plants and (depending on the pollination treatment) had either a negative effect or no effect on daily display size. Conclusions. We found evidence that within-population variation in floral longevity in M. guttatus can be caused by correlations with flower size and can have consequences for daily display size that vary depending on the pollination environment. These results contrast with among-species studies of variation in floral longevity, which have not included display size but have found that longevity either is extended in species with larger flowers or is not related to flower size. Consequently, predictions about the causes of within-population variation in floral longevity based on among-species studies may be misleading. [ABSTRACT FROM AUTHOR]

Sexual dimorphism is expressed as different morphologies between the sexes of a species. Dimorphism is pronounced in gynodioecious populations which consist of female and hermaphrodite individuals. The small size of female flowers in gynodioecious species is often explained by resource re‐allocation to seed production instead of large flowers. However, pollinator attraction is critical to female fitness, and factors other than resource savings are needed to explain the small size of female flowers. We hypothesized that the floral size dimorphism in the perennial gynodioecious Geranium sylvaticum (L.) is adaptive in terms of pollination. To test this "pollination hypothesis," we video recorded the small female and large hermaphrodite G. sylvaticum flowers. We parameterized floral visitor behavior when visiting a flower and calculated pollination probabilities by a floral visitor as the probability of touching anther and stigma with the same body part. Pollination probability differed in terms of flower sex and pollinator species. Bumblebees had the highest pollination probability. The small female flowers were more likely to receive pollen via several pollinator groups than the large hermaphrodite flowers. The pollen display of hermaphrodites matched poorly with the stigma display of hermaphrodites, but well with that of females. Although the small size of female flowers is commonly explained by resource re‐allocation, we show that sexual dimorphism in flower size may increase the main reproductive functions of the females and hermaphrodites. Dimorphism increases pollination probability in females and fathering probability of the hermaphrodites likely driving G. sylvaticum populations towards dioecy. [ABSTRACT FROM AUTHOR]

Premise: Ex situ cultivation is important for plant conservation, but cultivation in small populations may result in genetic changes by drift, inbreeding, or unconscious selection. Repeated inbreeding potentially influences not only plant fitness, but also floral traits and interactions with pollinators, which has not yet been studied in an ex situ context. Methods: We studied the molecular genetic variation of Digitalis lutea from a botanic garden population cultivated for 30 years, a frozen seed bank conserving the original genetic structure, and two current wild populations including the source population. In a common garden, we studied the effects of experimental inbreeding and between‐population crosses on performance, reproductive traits, and flower visitation of plants from the garden and a wild population. Results: Significant genetic differentiation was found between the garden population and the wild population from which the seeds had originally been gathered. After experimental selfing, inbreeding depression was only found for germination and leaf size of plants from the wild population, indicating a history of inbreeding in the smaller garden population. Moreover, garden plants flowered earlier and had floral traits related to selfing, whereas wild plants had traits related to attracting pollinators. Bumblebees visited more flowers of outbred than inbred plants and of wild than garden plants. Conclusions: Our case study suggests that high levels of inbreeding during ex situ cultivation can influence reproductive traits and thus interactions with pollinators. Together with the effects of genetic erosion and unconscious selection, these changes may affect the success of reintroductions into natural habitats. [ABSTRACT FROM AUTHOR]

We exploited ecotypic variation in Impatiens capensis (Balsaminaceae) to test the hypothesis that mutation accumulation accompanying relaxed selection on the chasmogamous (CH) flower leads to more variable flower shapes and smaller flowers. Sun ecotype populations of this species occur along sunny riverbanks and marshes and produce both CH and cleistogamous (CL) flowers, while shade ecotype populations occur in shady forest floors and produce only CL flowers. In the shade ecotype, it is assumed that selection on the CH flower has been relaxed. Seedlings from population samples of the two ecotypes exhibited different first internode growth responses to low ratios of red:far-red light as seen in earlier studies of ecotypic differentiation to sunny versus shady conditions, helping to verify historical illumination conditions in the populations. We examined the CH floral mutation accumulation hypothesis by comparing the shape and size of the modified CH flower sepal under greenhouse conditions that triggered CH flowering in plants of both ecotypes. Contrary to our predictions, geometric morphometric analysis of sepal shape variation indicated little difference between the shade and sun ecotypes. We suggest that mutations that influence CH flower shape may have pleiotropic effects on structures or processes that remain under selection even when the CH flowers are not produced. On the other hand, sepal size was significantly smaller in the shade ecotype populations. In the case of CH sepal size, the mutational effects appear to be directional, towards producing smaller sepals, and would likely be deleterious if shade ecotype plants encounter sunnier conditions where the plants can produce CH flowers. [ABSTRACT FROM AUTHOR]

Geographic differences in floral traits may reflect geographic differences in effective pollinator assemblages. Independent local adaptation to pollinator assemblages in multiple regions would be expected to cause parallel floral trait evolution, although sufficient evidence for this is still lacking. Knowing the intraspecific evolutionary history of floral traits will reveal events that occur in the early stages of trait diversification. In this study, we investigated the relationship between flower spur length and pollinator size in 16 populations of Aquilegia buergeriana var. buergeriana distributed in four mountain regions in the Japanese Alps. We also examined the genetic relationship between yellow‐ and red‐flowered individuals, to see if color differences caused genetic differentiation by pollinator isolation. Genetic relationships among 16 populations were analyzed based on genome‐wide single‐nucleotide polymorphisms. Even among populations within the same mountain region, pollinator size varied widely, and the average spur length of A. buergeriana var. buergeriana in each population was strongly related to the average visitor size of that population. Genetic relatedness between populations was not related to the similarity of spur length between populations; rather, it was related to the geographic proximity of populations in each mountain region. Our results indicate that spur length in each population evolved independently of the population genetic structure but in parallel in response to local flower visitor size in different mountain regions. Further, yellow‐ and red‐flowered individuals of A. buergeriana var. buergeriana were not genetically differentiated. Unlike other Aquilegia species in Europe and America visited by hummingbirds and hawkmoths, the Japanese Aquilegia species is consistently visited by bumblebees. As a result, genetic isolation by flower color may not have occurred. [ABSTRACT FROM AUTHOR]

In many communication systems, signal receivers profit from honest signals that indicate the signaller's quality, whereas low‐quality signallers should profit from cheating. Under such a conflict of interests between signallers and signal receivers, the maintenance of honest signals presents a puzzle. In theory, honesty can represent an evolutionarily stable strategy, but the actual mechanisms have been studied in few systems only.Here, we investigate honest signalling in a plant species, Brassica rapa, that advertises nectar volumes to pollinators by two honest floral signals; corolla size and the floral volatile phenylacetaldehyde. In a series of experiments we tested for physiological constraints and pollinator behaviours related to honest floral signals and nectar volume, incorporated the result into a mathematical model, and verified its predictions experimentally.While honest floral signals attracted pollinators, the bees' flower visitation time depended on nectar volume and was associated with the number of seeds that flowers developed. Furthermore, honest floral signals and seed set without pollen limitation both increased after soil fertilisation, indicating nutrient limitation in these traits.The mathematical model which incorporates these findings showed that honest signalling in B. rapa can be maintained by a combination of pollinator behaviour and resource limitation causing differential benefits of nectar production. The study demonstrates how honest floral signalling can evolve as a stable strategy in a plant species. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Background: Geographic differences in floral size sometimes reflect geographic differences in pollinator size. However, we know little about whether this floral size specialization to the regional pollinator size occurred independently at many places or occurred once and then spread across the distribution range of the plant species. Results: We investigated the relationship between the local floral size of flowers and local pollinator size in 12 populations of Lamium album var. barbatum on two different mountains in the Japan Alps. Then, using 10 microsatellite markers, we analyzed genetic differentiation among the 12 populations. The results showed that local floral size was correlated with the average size of relevant morphological traits of the local pollinators: floral size was greater in populations visited frequently by the largest flower visitors, Bombus consobrinus queens, than it was in other populations. We also found that the degree of genetic similarity between populations more closely reflected interpopulation geographic proximity than interpopulation similarity in floral size. Conclusions: Although genetic similarity of populations was highly associated with geographic proximity, floral size varied independently of geographic proximity and was associated with local pollinator size. These results suggest that in L. album var. barbatum, large floral size evolved independently in populations on different mountains as a convergent adaptation to locally abundant large bumblebee species. [ABSTRACT FROM AUTHOR]

The high variability observed in floral traits has been interpreted as resulting from the adaptation of plants to pollinators, as the latter present innate preferences for specific floral traits and impose selection over them. However, some pollinators such as bees can learn to associate floral signals with rewards, thus increasing floral constancy on more rewarding flowers. The integration of all these rewards and cues is markedly important at community level, where co‐flowering species compete for pollinators.In order to verify the honesty of the above‐mentioned signals, we examined the association between floral visual signals (size, colour, symmetry and floral display) and rewards (pollen and nectar) for 98 species in a Mediterranean community. The associations between floral traits were analysed considering the phylogenetic relationship between the different species.Flower colour, size, pollen volume or amount of sucrose exhibited no phylogenetic signal, which suggests an adaptive evolution in response to different conditions in the pollinator community. Flower size was seen to constitute the most honest signal for pollinators, as this was significantly associated with quantities of pollen and nectar. In contrast, nectar concentration was observed to be positively associated with chromatic contrast. We detected no relationship between flower shape and rewards, on the one hand, nor between flower display and rewards, on the other.Our study unequivocally demonstrates the correlation between rewards and the visual signals perceived by bees, the most effective pollinators in the Mediterranean Basin. In the community studied, bees employed flower size at longer distances and chromatic contrast at shorter distances to predict rewards. The limited number of studies existing in this sense indicates that this kind of association appears to be community specific. [ABSTRACT FROM AUTHOR]

Background and Aims Sexual dimorphism for floral traits is common in dioecious plant species. Beyond its significance for understanding how selection acts on plant traits through male vs. female reproductive function, sexual dimorphism has also been proposed as a possible risky characteristic for insect-pollinated plants, as it could drive pollinators to forage mostly on male plants. However, even though most flowering plant species spread their flowering across several weeks or months, the temporal variation of floral phenotypes and sexual dimorphism have rarely been investigated. Methods We performed a survey of male and female plants from the dioecious generalist-pollinated Silene dioica (Caryophyllaceae) in a common garden experiment, over two consecutive flowering seasons. Flower number and floral size were measured each week, as well as pollen quantity and viability in male plants. Key Results Sexual dimorphism was found for all investigated floral traits, with males showing an overall higher investment in flower production and flower size. Males and females showed a similar temporal decline in flower size. The temporal dynamics of daily flower number differed between sexes, with males showing a peak in the middle of their flowering season, whereas flower production by females was quite stable over time. At the scale of the experimental population, both individual and floral sex ratios appeared to vary across the flowering season. Moreover, because the onset of flowering varied among plants, the magnitude of sexual dimorphism in floral size also fluctuated strongly through time. Conclusions Capturing male/female differences with only one temporal measurement per population may not be informative. This opens stimulating questions about how pollinator behaviour and resulting pollination efficiency may vary across the flowering season. [ABSTRACT FROM AUTHOR]

Theoretical models indicate that the structure of plant–pollinator networks has important implications for the reproduction and survival of species. However, despite the growing information on the mechanisms underlying such a structure, it is still difficult to predict the functional consequences of species' structural positions in these networks. From the plant perspective, species position and roles in pollination networks might be related to traits describing flower attractiveness, availability and dependence on pollinators. In turn, both network metrics and plant traits might influence plant species fitness.During two field seasons, we collected data from the 23 most abundant plant species in a rich coastal community, to evaluate the association between population and floral traits (floral abundance at population level and flowers per individual, flower shape and size, flowering length, nectar volume, pollinator dependence), species‐level network metrics (linkage level, specialization –d′–, weighted closeness centrality, network roles related to modularity) and plant fitness (seeds/flower, seed weight).Flowering length, flower size, flower abundance and pollinator dependence were positively related to greater generalization, as measured by various indices. More abundant species and those with larger flowers showed higher linkage levels (i.e. higher number of pollinator species), whereas longer flowering periods were negatively related to d′ and positively related to closeness centrality and important roles in the network. Likewise, plants species more dependent on pollinators occupied more central positions in the network. Furthermore, centrality in the networks was significantly associated with plant fitness. Specifically, central species in the network produced more and heavier seeds than the others. However, other plant traits, such as flower size and pollinator dependence had additional direct effects on seed production.Synthesis. Our study highlights how population and floral traits define the positions and roles of species structuring the pollination communities. Moreover, the relationships between network metrics and plant reproduction indicate, for the first time, the functional implications of these structural positions at the interspecific level of community assembly. [ABSTRACT FROM AUTHOR]

Sexual dimorphism is a pervasive form of variation within species. Understanding how and why sexual dimorphism evolves would contribute to elucidating the mechanisms underlying the diversification of traits. In flowering plants, pollinators are considered a driver of sexual dimorphism when they affect female and male plant fitness in distinct ways. Here, we found that flowers appear to manipulate the behavior of pollinators using sexually dimorphic traits in the dioecious tree Eurya japonica. In this plant, female flowers present a higher-quality reward for pollinators, whereas male flowers have a more conspicuous appearance. Plants benefit by inducing pollinators to carry pollen from male to female flowers, and their sexual dimorphism might thus facilitate pollen movement through pollinator behavior. In two-choice experiments, pollinators frequently moved from male to female flowers, whereas computer simulation suggested that sexually dimorphic traits would evolve if pollinators changed behavior depending on the traits of the flowers they had just visited. These results suggest that the floral traits affecting the visiting order of pollinators have evolved in plants. Using E. japonica, we theoretically show that the induction of sequential behavior in pollinators might be crucial to the evolution of sexual dimorphism in flowers, and our experiments support these findings. [ABSTRACT FROM AUTHOR]

Premise: Understanding how environmental stress affects the strength of mutualisms is critically important given observed and projected environmental changes. In particular, the frequency and duration of drought have been increasing worldwide. We investigated how water availability affects plant traits that mediate a pollination mutualism. Methods: For butterfly‐pollinated Phlox drummondii, we determined how moisture availability affects flower size, nectar volume, and nectar sugar amount. Furthermore, we explored the role that local adaptation may play in responses to moisture availability by including plants collected from regions that differ in aridity. Finally, we determined whether responses of plant populations to selection may differ under drought by calculating heritability of traits under control and dry conditions. Results: Flower size was generally smaller in dry plants than in control plants. Early in the treatment period, nectar volume and sugar were higher in dry plants than in control plants for plants from both arid and wetter regions. With prolonged dry treatment, nectar volume and sugar remained higher only in plants from the arid region. Heritability of floral traits was lower for water‐limited plants than for control plants. Conclusions: Plant investment into pollination mutualisms under environmental stress may depend on the extent to which populations are already locally adapted to such conditions, suggesting that mutualism may remain strong, at least in arid regions. However, decreases in heritability under water‐limitation suggest that responses to selection imposed by pollinators may be low, even if drought‐adapted plants maintain production of rewards to pollinators. [ABSTRACT FROM AUTHOR]

For most angiosperms, flowers are critical to reproduction because they increase rates of outcrossing. Flowers are highly variable in numerous traits, including size, shape, and color. Most of this variation is thought to have arisen because of selection by pollinators. Yet nonpollinator selection is increasingly being recognized as contributing to floral trait evolution. Nonpollinator agents of selection that often oppose pollinator selection include the physiological and resource costs of producing and maintaining flowers. Here, I (1) summarize recent studies on the macroevolution of floral physiological traits and (2) apply an energy balance model to examine how two pollination traits (flower color and flower size) can interact with hydraulic traits to influence flower physiology. These modeling results show that under certain conditions flower color variation can overwhelm the effects of floral transpiration and flower size variation on flower temperature. Using a novel data set of flower size from the California flora, I show that the range of flower size most common in the California flora is the range in which complex, nonlinear dynamics in flower energy balance occur. These results suggest that floral traits under selection by biotic agents can have large implications for flower physiology, and only some of these potentially deleterious effects can be offset by flower hydraulic traits. These complex interactions between pollination traits (flower size and color) and physiological traits (surface conductance to water vapor) suggest that a more unified framework for understanding the evolution of floral form and function would simultaneously consider the interaction between physiological traits and traits under biotic selection. [ABSTRACT FROM AUTHOR]

In angiosperms, a decrease in fruit production towards the apex of individual inflorescences is usually observed. Orchids are thought to be primarily pollination‐limited species, and non‐uniform pollination could cause this decrease pattern in several species. Fruit production was investigated in relation to flower position and floral display size in Orchis militaris (Orchidaceae), a deceptive species.Over 2 years, eight populations of O. militaris were studied and fruit position along the inflorescence was recorded. Generalised linear models were performed to examine the effect of population, year, flower position and floral display size on fruit production.The dominant pattern was characterised by a higher fruit set in the middle part of the inflorescence (parabolic pattern). A non‐directional pattern of fruit production was also detected in some populations. Within a given population, patterns were generally consistent among years. In one of the two study years and in one of the eight populations specifically, the proximal‐to‐distal decrease in fruit production was dramatic in plants with a large floral display but weak or absent in small displays.Our study demonstrates the intraspecific diversity of fruit distribution patterns in O. militaris. Non‐uniform pollination along the inflorescence is likely to be responsible for the parabolic pattern, while irregular visitation could explain the non‐directional pattern of fruit production. Pattern variation among years and between populations could arise from spatiotemporal variation in pollinator assemblages. Resource competition effects could explain the interaction effect between display size and flower position. [ABSTRACT FROM AUTHOR]

Premise of research. Variation in floral traits is mainly attributed to variations in the abundance or taxonomic composition of the pollinator assemblage. The diversity and abundance of insect pollinators decrease with elevation, which might modify the allocation patterns of floral traits in plants (i.e., attraction or rewards) at high elevation to compete for pollinators. This competition would be pronounced between congeneric and sympatric species. In this study, we evaluated the pattern of flower size variation and the composition of floral visitor assemblages along elevation in two congeneric and sympatric species of Anthurium. Methodology. We investigated the variation of floral traits and the variation in the flower-visitor assemblages in Anthurium caucanum and Anthurium panduriforme in seven transects located every 100 m of elevation along an elevation gradient (2200–2800 m a.s.l.) for 1 yr. We measured floral size (spadix length and diameter and spathe length and width) and functional traits (number of flowers, ovules, and pollen grains; stigmatic area; and pollen-to-ovule ratio) in plants from each species. Pivotal results. Spadix length increased with elevation in A. panduriforme , while it decreased in A. caucanum. Pollen number (i.e., male function) and pollen-to-ovule ratio were higher in A. panduriforme compared with A. caucanum , while the opposite occurred with stigmatic area (i.e., female function). The abundance of flower visitors decreased at high elevation. Conclusions. The contrasting patterns of floral size variation, allocation patterns to the reproductive male and female functions, and pollen-to-ovule ratios suggest that the variation in the pollination context along elevation seems to have promoted differences in inflorescence morphology among these congeneric and co-occurring species, most likely to reduce competition for pollinators. This could drive the species to different reproductive and mating strategies, allowing them to be successful in the shared environment. [ABSTRACT FROM AUTHOR]

Animals that rely on nectar are expected to display floral trait preferences correlating to the signals of nectar source flowers. Batesian mimicry evolves to exploit these pre-existing signal–receiver relationships, attracting pollinators through an adaptive resemblance to specific co-occurring rewarding species. The nectar-feeding long-proboscid flies of South Africa are pollinators for several deceptive orchid species that are putatively Batesian mimics. We tested whether flies' measured color preference varied among communities providing different nectar-source diets, which would indicate the necessary signal–receiver conditions for the evolution of advergent Batesian mimicry. We introduced artificial rewardless flowers into flowering communities that supported divergent nectar diets in resident flies and inferred floral trait preferences of a long-proboscid fly species (Prosoeca ganglbaueri) from visitation behavior to these artificial flowers. The experiment showed that the preference of flies for white versus pink was strongly predicted by the colors of flowers most commonly visited by flies at a site. Furthermore, generalization in preference was positively correlated with the variance in nectar-community hue, i.e. flies showed more generalized preference in more spectrally diverse flower communities. The floral tube length of local nectar sources also influenced how readily the flies probed the artificial flowers during attempted foraging. These results support the hypothesis that nectarless orchids pollinated by P. ganglbaueri experience selection for traits that exploit site-specific mutualistic relationships between fly pollinators and their local floral communities. [ABSTRACT FROM AUTHOR]

Pollen limitation can strongly influence reproduction of pollinator-dependent plants. Flower abundance can affect pollination ‘quantity’ and ‘quality’ due to its influence on pollen availability and foraging patterns of pollinators, ultimately impacting on seed production. We complemented individual-based measurements with landscape-level metrics to assess the influence of conspecific flower availability at different spatial scales on the quantity and quality components of pollination, and their impact on seed production in the invasive shrub Cytisus scoparius. In 2013-2014, we sampled 40 C. scoparius populations in Nahuel Huapi National Park, Patagonia (Argentina). In each population, we estimated the proportion of tripped flowers, fruit- and seed-set in five randomly selected individuals. The proportion of tripped flowers and the proportion of them setting fruit were used as proxies of pollination quantity and quality, respectively. Conspecific flower availability at distinct spatial scales (5-1000 m) was estimated as the area covered by flowering C. scoparius from color aerial photographs. Flower availability influenced seed output due to contrasting scale-dependent effects on pollination quantity and quality. Increasing flower availability at the landscape-scale reduced pollination quantity, whereas at the neighborhood-scale it increased pollination quality. The overall positive effect of flower availability on seed output at the neighborhood scale was slightly higher than the overall negative effect at the landscape scale. Moreover, pollination quality had a higher positive effect on seed output than pollination quantity. Our results demonstrate that pollination quality may severely limit plant reproduction. Pollination quality limitation can act independently of pollination quantity limitation because these factors operate at different spatial scales. [ABSTRACT FROM AUTHOR]

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Flowers represent a key innovation during plant evolution. Driven by reproductive optimization, evolution of flower morphology has been central in boosting species diversification. In most cases, this has happened through specialized interactions with animal pollinators and subsequent reduction of gene flow between specialized morphs. While radiation has led to an enormous variability in flower forms and sizes, recurrent evolutionary patterns can be observed. Here, we discuss the targets of selection involved in major trends of pollinator-driven flower evolution. We review recent findings on their adaptive values, developmental grounds and genetic bases, in an attempt to better understand the repeated nature of pollinator-driven flower evolution. This analysis highlights how structural innovation can provide flexibility in phenotypic evolution, adaptation and speciation. [ABSTRACT FROM AUTHOR]

There is discussion over whether pollen limitation exerts selection on floral traits to increase floral display or selects for traits that promote autonomous self-fertilization. Some studies have indicated that pollen limitation does not mediate selection on traits associated with either pollinator attraction or self-fertilization. Primula tibetica is an inconspicuous cross-fertilized plant that may suffer from pollen limitation. We conducted a selection analysis on P. tibetica to investigate whether pollen limitation results in selection for an increased floral display in case the evolution of autonomous self-fertilization has been difficult for this plant. The self- and intra-morph incompatibility features, the capacity for autonomous self-fertilization, and the magnitude of pollen limitation were examined through hand-pollination experiments. In 2016, we applied selection analysis on the flowering time, corolla width, stalk height, flower tube length, and flower number in P. tibetica by tagging 76 open-pollinated plants and 37 hand-pollinated plants in the field. Our results demonstrated that P. tibetica was strictly self- and intra-morph incompatible. Moreover, the study population underwent severe pollen limitation during the 2016 flowering season. The selection gradients were found to be significantly positive for flowering time, flower number, and corolla width, and marginally significant for the stalk height. Pollinator-mediated selection was found to be significant on the flower number and corolla width, and marginally significant on stalk height. Our results indicate that the increased floral display may be a vital strategy for small distylous species that have faced difficulty in evolving autonomous self-fertilization. [ABSTRACT FROM AUTHOR]

Background: Flower size varies dramatically across angiosperms, representing innovations over the course of >130 million years of evolution and contributing substantially to relationships with pollinators. However, the genetic underpinning of flower size is not well understood. Saltugilia (Polemoniaceae) provides an excellent nonmodel system for extending the genetic study of flower size to interspecific differences that coincide with variation in pollinators. Results: Using targeted gene capture methods, we infer phylogenetic relationships among all members of Saltugilia to provide a framework for investigating the genetic control of flower size differences via RNA-Seq de novo assembly. Nuclear concatenation and species tree inference methods provide congruent topologies. The inferred evolutionary trajectory of flower size is from small flowers to larger flowers. We identified 4 to 10,368 transcripts that are differentially expressed during flower development, with many unigenes associated with cell wall modification and components of the auxin and gibberellin pathways. Conclusions: Saltugilia is an excellent model for investigating covarying floral and pollinator evolution. Four candidate genes from model systems (BIG BROTHER, BIG PETAL, GASA, and LONGIFOLIA) show differential expression during development of flowers in Saltugilia, and four other genes (FLOWERING-PROMOTING FACTOR 1, PECTINESTERASE, POLYGALACTURONASE, and SUCROSE SYNTHASE) fit into hypothesized organ size pathways. Together, these gene sets provide a strong foundation for future functional studies to determine their roles in specifying interspecific differences in flower size. [ABSTRACT FROM AUTHOR]

Animals, including human beings, tend to respond more strongly to stimuli that are associated with the highest relative rewards. This applies not only to food rewards but also to reproductive success. In the present review article this issue is discussed for insects in connection with intersexual communication and flower-visiting behaviour. Implications of the preference for supernormal visual releasing stimuli are examined from a sensory and evolutionary perspective, including a consideration of the choice of potential mates and recognition of the most rewarding flowers. [ABSTRACT FROM AUTHOR]

Flower size is a key trait in the reproductive ecology of animal-pollinated plants. However, pollinator-mediated selection does not always modulate this trait and environmental conditions and/or antagonist interactions may favor smaller flowers. We evaluate the occurrence of a large-flowered family in a hot and dry Mediterranean environment, mediated by a cost-benefit balance and a male-female conflict. Large flowers have sizeable benefits in terms of pollination and reproductive success and pollinators mediate selection through male function, but female fitness is context-dependent. High floral production and maintenance costs and florivore incidence in large flowers limit female function, which counteracts pollinator-mediated selection. Large flowers are highly costly in the Mediterranean and flower size is mediated by a sexual conflict between the benefits of male function and the costs of the female one. However, a short floral longevity, occasional pollen limitation and selection through maleness keep the existence of large flowers in these environments. [ABSTRACT FROM AUTHOR]

Background: How floral traits and community composition influence plant specialization is poorly understood and the existing evidence is restricted to regions where plant diversity is low. Here, we assessed whether plant specialization varied among four species-rich subalpine/alpine communities on the Yulong Mountain, SW China (elevation from 2725 to 3910 m). We analyzed two factors (floral traits and pollen vector community composition: richness and density) to determine the degree of plant specialization across 101 plant species in all four communities. Floral visitors were collected and pollen load analyses were conducted to identify and define pollen vectors. Plant specialization of each species was described by using both pollen vector diversity (Shannon's diversity index) and plant selectiveness (d' index), which reflected how selective a given species was relative to available pollen vectors. Results: Pollen vector diversity tended to be higher in communities at lower elevations, while plant selectiveness was significantly lower in a community with the highest proportion of unspecialized flowers (open flowers and clusters of flowers in open inflorescences). In particular, we found that plant species with large and unspecialized flowers attracted a greater diversity of pollen vectors and showed higher selectiveness in their use of pollen vectors. Plant species with large floral displays and high flower abundance were more selective in their exploitation of pollen vectors. Moreover, there was a negative relationship between plant selectiveness and pollen vector density. Conclusions: These findings suggest that flower shape and flower size can increase pollen vector diversity but they also increased plant selectiveness. This indicated that those floral traits that were more attractive to insects increased the diversity of pollen vectors to plants while decreasing overlap among co-blooming plant species for the same pollen vectors. Furthermore, floral traits had a more important impact on the diversity of pollen vectors than the composition of anthophilous insect communities. Plant selectiveness of pollen vectors was strongly influenced by both floral traits and insect community composition. These findings provide a basis for a better understanding of how floral traits and community context shape interactions between flowers and their pollen vectors in species-rich communities. [ABSTRACT FROM AUTHOR]

The astonishing diversity of floral form in angiosperm plants is driven in large part by preferences of pollinators for various floral traits, including learned preferences. Remarkably, almost all of a vast literature on learning and memory in pollinators relates to nectar as a reward, even though bees and many flies, beetles, and butterflies must collect pollen. In this study, we asked if bees formed preferences for plant species from which pollen had been collected successfully. Using absolute conditioning, we gave pollen foraging bees experience with plant species that offered only pollen rewards. Naive bees generally showed modest preferences, whereas experienced bees adopted strong preferences for those species over alternative species not previously experienced. Learned preferences were retained for at least 24 h, consistent with preferences learned with nectar rewards. These experience-mediated changes in preference raised the possibility that bees formed associations between particular floral features and pollen rewards. We therefore asked if learned preferences required that bees successfully collect pollen. Using differential conditioning, we determined that learned preferences were strongly influenced by receipt of a pollen reward. In a final experiment, we characterized the importance of 2 floral features, the corolla and the anther, in the expression of learned preferences. Although experience altered responses to both floral parts, responses to anthers were influenced more strongly. We discuss recent evidence in the literature for associative learning with pollen rewards and propose that learned preferences in the context of pollen collection have played an important role in floral display evolution. [ABSTRACT FROM AUTHOR]

The effects of climate change on species interactions are poorly understood. Investigating the mechanisms by which species interactions may shift under altered environmental conditions will help form a more predictive understanding of such shifts. In particular, components of climate change have the potential to strongly influence floral volatile organic compounds ( VOCs) and, in turn, plant-pollinator interactions. In this study, we experimentally manipulated drought and herbivory for four forb species to determine effects of these treatments and their interactions on (1) visual plant traits traditionally associated with pollinator attraction, (2) floral VOCs, and (3) the visitation rates and community composition of pollinators. For all forbs tested, experimental drought universally reduced flower size and floral display, but there were species-specific effects of drought on volatile emissions per flower, the composition of compounds produced, and subsequent pollinator visitation rates. Moreover, the community of pollinating visitors was influenced by drought across forb species (i.e. some pollinator species were deterred by drought while others were attracted). Together, these results indicate that VOCs may provide more nuanced information to potential floral visitors and may be relatively more important than visual traits for pollinator attraction, particularly under shifting environmental conditions. [ABSTRACT FROM AUTHOR]

The concept of pollinator niche complementarity maintains that species-rich pollinator communities can provide higher and more stable pollination services than species-poor communities, due to contrasting spatial and/or temporal pollination activity among groups of pollinators. Complementarity has usually been examined in pollinators' patterns of flower visitation or abundance, while largely neglecting the possibility of complementarity in patterns of single-visit contribution to fruit/seed set (pollination efficiency). However, variability in pollination efficiency can greatly affect pollinators' overall pollination services and may therefore contribute an additional, important aspect of complementarity., In this study, we investigated the existence of pollinator complementarity in both visitation rates and pollination efficiencies. The study was conducted in 43 watermelon fields cultivated for seed consumption in a Mediterranean agro-natural landscape in central Israel. We studied spatiotemporal variation in pollinators' visitation activity, measured by repeated observations and netting, and single-visit pollination efficiency, measured by the fruit and seed set rates of hermaphrodite flowers exposed to a single bee visit. Visitation and pollination efficiency were measured throughout the day and season, within and between fields with contrasting availability of nearby wild plants, and among flowers of different sizes., Pollinator species' visitation rates as well as single-visit fruit set efficiencies, but not seed set efficiencies, exhibited significant spatiotemporal variation that contributed to their complementarity. Pollinators' visit frequencies were affected by surrounding land use, location within field, time throughout the season, and time of day. Pollinators' fruit set efficiencies were affected by ovary size and time of day., Synthesis and applications. Crop pollinators may exhibit complementarity in both their visitation rates and pollination efficiencies, which can promote the overall level and stability of their pollination services. Complementarity in pollination efficiencies suggests further diversity effects on crop yield, and calls for taking into account the variability in pollination efficiency along spatiotemporal scales rather than considering it a constant, species-specific trait. However, some modes of niche complementarity may not necessarily translate into increased pollination services and crop yield; the relevance and limitations of such mechanisms should be considered in the light of the specific crop and management system studied. [ABSTRACT FROM AUTHOR]

Background and Aims Flowering plants display considerable variation in mating system, specifically the relative frequency of cross- and self-fertilization. The majority of estimates of outcrossing rate do not account for temporal variation, particularly during the flowering season. Here, we investigated seasonal variation in mating and fertility in Incarvillea sinensis (Bignoniaceae), an annual with showy, insect-pollinated, 'one-day' flowers capable of delayed selfing. We examined the influence of several biotic and abiotic environmental factors on day-to-day variation in fruit set, seed set and patterns of mating. Methods We recorded daily flower number and pollinator abundance in nine 33-m patches in a population at Mu Us Sand land, Inner Mongolia, China. From marked flowers we collected data on daily fruit and seed set and estimated outcrossing rate and biparental inbreeding using six microsatellite loci and 172 open-pollinated families throughout the flowering period. Key Results Flower density increased significantly over most of the 50-d flowering season, but was associated with a decline in levels of pollinator service by bees, particularly on windy days. Fruit and seed set declined over time, especially during the latter third of the flowering period. Multilocus estimates of outcrossing rate were obtained using two methods (the programs MLTR and BORICE) and both indicated high selfing rates of 80%. There was evidence for a significant increase in levels of selfing as the flowering season progressed and pollinator visitation declined. Biparental inbreeding also declined significantly as the flowering season progressed. Conclusions Temporal variation in outcrossing rates may be a common feature of the mating biology of annual, insect-pollinated plants of harsh environments but our study is the first to examine seasonal mating-system dynamics in this context. Despite having large flowers and showy floral displays, I. sinensis attracted relatively few pollinators. Delayed selfing by corolla dragging largely explains the occurrence of mixed mating in I. sinensis, and this mode of self-fertilization probably functions to promote reproductive assurance when pollinator service is limited by windy environmental conditions and competition from co-occurring flowering plants. [ABSTRACT FROM AUTHOR]

Studies of factors influencing spatial variation in flower size offer insights into floral evolution. We investigated altitudinal variations in five flower dimensions of two native Japanese Impatiens species (I. textori and I. noli-tangere) and their interactions with their faunal visitors. These two species have similar floral traits, including flower shape, flowering time, and pollinator species; both species are pollinated mainly by Bombus diversus. In I. textori, all measured flower dimensions were negatively correlated with altitude. In contrast, in I. noli-tangere, no measured flower dimensions correlated with altitude. Thus, the altitudinal pattern of flower size variation differed between these congeneric co-habiting herbaceous plant species. The different patterns suggest that the factors (e.g. altitudinal variations of abiotic factors) that cause variation of flower size differ between these two Impatiens species even though focal species have similar floral traits (e.g. flower shape, flowering time, and pollinator species). [ABSTRACT FROM AUTHOR]

The pollen/ovule (P/O) ratio has been used as predictor of the reproductive system of angiosperms (lowest P/O values correspond to obligate autogamous species while the highest correlate with obligate xenogamous species) but it does not keep for all the taxa. The relation of P/O index with pollen and stigma size, flower diameter, pollen vitality, stigmatic area/pollen-bearing area of the pollinators ratio (SA/PBA), taxonomy and breeding system was analyzed in four Passiflora species with different degree of compatibility, pollination system and taxonomic placement. P/O of the self-compatible species (with shorter anthesis, scarce pollinator visits or minor longevity) was lower than P/O of the self-incompatible ones. The P/O values of the outcrossers could be related with the highly efficient pollination performed by the frequently visiting Xylocopa species; these bees transport Passiflora pollen in a usually monospecific mass on the thorax, resembling grouped pollen dispersal. All the taxa exhibited pollen with unimodal size and high vitality, except for P. misera, in which pollen size was bimodal, with similar quantities of grains in both classes, the large being more vital than the small ones. Correlations of P/O were positive with flower size, negative with stigma area/pollen-bearing area of the pollinator, null with stigma area and not significant with pollen size. The P/O ratio did not reflect taxonomic affinities; this index and the breeding system were more related with pollinator type, anthesis, flower size and SA/PBA than with pollen grain or stigma size.

PREMISE OF THE STUDY: The evolutionary shift from radial to bilateral symmetry in flowers is generally associated with the evolution of low flower-size variation. This phenomenon supports the hypothesis that the lower size variation in bilateral flowers can be attributed to low pollinator diversity. In this study, we propose two other hypotheses to explain low flower-size variation in bilateral symmetrical flowers. To test the three hypotheses, we examined the relative importance of pollinator diversity, composition, and bilateral symmetry itself as selective forces on low flower-size variation. METHODS: We examined pollinator diversity and composition and flower-size variation for 36 species in a seminatural ecosystem with high bee richness and frequent lepidopteran visitation. KEY RESULTS: Bilateral flowers were more frequently visited than radial flowers by larger bees, but functional-group diversity of the pollinators did not differ between symmetry types. Although bilateral flowers had significantly lower flower-size variation than radial flowers, flower-size variation did not vary with pollinator diversity and composition but was instead related to bilateral symmetry. CONCLUSIONS: Our results suggest that the lower size variation in bilateral flowers might have evolved under selection favoring the control of pollinator behavior on flowers to enhance the accurate placement of pollen on the body of the pollinator, independent of pollinator type. Because of the limited research on this issue, future work should be conducted in various types of plant-pollinator communities worldwide to further clarify the issue. [ABSTRACT FROM AUTHOR]

Premise of research. The size of floral display, flower, and reward can influence pollinator foraging behavior. However, the importance of correlations between floral traits and rewards in affecting pollinator preferences is unclear. Moreover, in monoecious plants and plants with strong within-flower dichogamy, some flowers do not offer a pollen reward, and pollinator attraction may depend on the number of rewarding flowers rather than floral display size. We examined these questions in the Rocky Mountain columbine, where bumblebees can forage only for pollen produced in male-phase flowers. Methodology. We used dual-choice assays and experimental arrays to determine whether a correlation between pollen reward and flower size was necessary before bees could distinguish between flowers with different pollen rewards or sizes. The impact of floral display size on bee foraging was first examined using inflorescences where all flowers provided a reward, either small or large. We then varied either floral display size or number of rewarding flowers while keeping the other variable constant. Pivotal results. Bumblebees preferred flowers with a larger pollen reward or larger flowers only when the size of the pollen reward was correlated with flower size. Bees selected larger floral displays when all flowers presented a similar amount of pollen but preferred inflorescences with more pollen-rewarding flowers when only a proportion of the flowers offered a pollen reward. Conclusions. Bees could identify whether a flower offered pollen and could quantify the number of pollenrewarding flowers. Correlations between floral traits and pollen reward affected pollinator preferences and may facilitate selection of floral traits by pollinators. Pollinators preferred inflorescences with more pollen-producing flowers rather than larger floral displays, which suggests that selection of floral display size by pollinators may differ between plants with different mating systems. [ABSTRACT FROM AUTHOR]

Stronger pollen limitation should increase competition among plants, leading to stronger selection on traits important for pollen receipt. The few explicit tests of this hypothesis, however, have provided conflicting support. Using the arithmetic relationship between these two quantities, we show that increased pollen limitation will automatically result in stronger selection (all else equal) although other factors can alter selection independently of pollen limitation. We then tested the hypothesis using two approaches. First, we analysed the published studies containing information on both pollen limitation and selection. Second, we explored how natural selection measured in one Ontario population of Lobelia cardinalis over 3 years and two Michigan populations in 1 year relates to pollen limitation. For the Ontario population, we also explored whether pollinator-mediated selection is related to pollen limitation. Consistent with the hypothesis, we found an overall positive relationship between selection strength and pollen limitation both among species and within L. cardinalis. Unexpectedly, this relationship was found even for vegetative traits among species, and was not found in L. cardinalis for pollinator-mediated selection on nearly all trait types. [ABSTRACT FROM AUTHOR]

The influence of locally different species interactions on trait evolution is a focus of recent evolutionary studies. However, few studies have demonstrated that geographically different pollinator-mediated selection influences geographic variation in floral traits, especially across a narrow geographic range. Here, we hypothesized that floral size variation in the Japanese herb Prunella vulgaris L. (Lamiaceae) is affected by geographically different pollinator sizes reflecting different pollinator assemblages. To evaluate this hypothesis, we posed two questions. (1) Is there a positive correlation between floral length and the proboscis length of pollinators (bumblebees) across altitude in a mountain range? (2) Does the flower-pollinator size match influence female and male plant fitness? We found geographic variation in the assemblage of pollinators of P. vulgaris along an altitudinal gradient, and, as a consequence, the mean pollinator proboscis length also changed altitudinally. The floral corolla length of P. vulgaris also varied along an altitudinal gradient, and this variation strongly correlated with the local pollinator size but did not correlate with altitude itself. Furthermore, we found that the size match between the floral corolla length and bee proboscis length affected female and male plant fitness and the optimal size match (associated with peak fitness) was similar for the female and male fitness. Collectively, these results suggest that pollinator-mediated selection influences spatial variation in the size of P. vulgaris flowers at a fine spatial scale. [ABSTRACT FROM AUTHOR]

Long corollas are a classical example of nectar barriers, because they prevent undesired visitors from consuming the reward intended for more effective pollinators. As the investment in nectar barriers increases, flower attractiveness and nectar rewards may also increase to maintain loyal visitation of most effective pollinators; and flowers may become more prone to nectar robbing. We evaluated the effect of nectar barriers (corolla tube length), two related traits (nectar volume and upper lip size) and the associated risk of nectar robbing, on the fecundity of Lonicera implexa plants from three populations differing in the abundance of its most efficient pollinator, the hummingbird hawkmoth Macroglossum stellatarum. Corolla tube length varied most among individuals within populations (45-46 % of total variance) and inflorescences within individuals (23-32 %), and showed little variation among populations (0.2-11 %). Longer corolla tubes were always associated with larger nectar volumes and larger upper lips, although the strength of the relationships varied across populations and years. Robbing frequency increased with corolla tube length, decreased with nectar volume and upper lip size, and its weak effects on fecundity were predominantly positive. Plant fecundity peaked at two different optima: long corollas with little nectar and short corollas with abundant nectar. However, the exact shape of the interaction between corolla length and nectar volume, as well as the combination of traits showing the highest fecundity, differed between populations and years. This variation could be explained by among-population differences in pollinator assemblages, and inter-annual changes in resources dedicated to reproduction. Our study shows that large nectar volumes can modulate the effect of corolla length as a nectar barrier, and that the combination of these two traits that maximises fecundity may be related to the identity of pollinators within each population. [ABSTRACT FROM AUTHOR]

In plants, the temporal pattern of floral displays, or display schedules, delimits an individual's mating opportunities. Thus, variation in the shape of display schedules can affect the degree of population synchrony and the strength of phenological assortative mating by flowering onset date. A good understanding of the mechanisms regulating the timing of flowering onset has been developed, but we know less about factors influencing subsequent patterns of floral display., We observed unusual multimodal display schedules in temperate populations of the annual legume Chamaecrista fasciculata. Here, we ask whether 'flowering pulses' are simultaneous among individuals and populations and explore potential underlying mechanisms and consequences of pulsed flowering., We monitored daily flower production for individual plants from genetically divergent populations during a series of field experiments that manipulated three potential influencers of display schedule shape: average daily temperature, pollinator availability and watering schedules. We measured floral longevity to isolate the contributions of flower retention and flower deployment to display schedules. We assessed relationships between flowering and environmental variables and compared estimates of population synchrony, individual synchrony and the strength of assortative mating with those of 29 unimodally flowering species from the area., We observed simultaneous flowering pulses in all experiments, with peaks aligned among individuals and populations despite variation in flowering onset and/or duration. Pulses were not the result of increases in average temperature, pollinator availability or variation in watering schedules. Seasonal fluctuations in temperature correlated with floral longevity and flower deployment, suggesting that the shape of display schedules may be plastic in response to fluctuations in temperature. Average population and individual synchrony differed only slightly from those of the species with unimodal schedules, while the average strength of assortative mating for flowering onset date was strongly reduced (0.21 in C. fasciculata vs. 0.35 for the 29 other species)., Synthesis. Researchers should take caution in assuming that components of display schedules are genetically or developmentally correlated with flowering onset. Variation in the shape of display schedules can influence patterns of gene flow within or between populations, with potential effects on the strength of phenological assortative mating and subsequent responses to selection. [ABSTRACT FROM AUTHOR]

Plant mating systems are driven by several pre-pollination factors, including pollinator availability, mate availability and reproductive traits. We investigated the relative contributions of these factors to pollination and to realized outcrossing rates in the patchily distributed mass-flowering shrub Rhododendron ferrugineum. We jointly monitored pollen limitation (comparing seed set from intact and pollen-supplemented flowers), reproductive traits (herkogamy, flower size and autofertility) and mating patterns (progeny array analysis) in 28 natural patches varying in the level of pollinator availability (flower visitation rates) and of mate availability (patch floral display estimated as the total number of inflorescences per patch). Our results showed that patch floral display was the strongest determinant of pollination and of the realized outcrossing rates in this mass-flowering species. We found an increase in pollen limitation and in outcrossing rates with increasing patch floral display. Reproductive traits were not significantly related to patch floral display, while autofertility was negatively correlated to outcrossing rates. These findings suggest that mate limitation, arising from high flower visitation rates in small plant patches, resulted in low pollen limitation and high selfing rates, while pollinator limitation, arising from low flower visitation rates in large plant patches, resulted in higher pollen limitation and outcrossing rates. Pollinator-mediated selfing and geitonogamy likely alleviates pollen limitation in the case of reduced mate availability, while reduced pollinator availability (intraspecific competition for pollinator services) may result in the maintenance of high outcrossing rates despite reduced seed production. [ABSTRACT FROM AUTHOR]

Pollinator-mediated selection is a major evolutionary driver of floral traits; yet, such selection has rarely been tested for floral extreme traits. The Oncocyclus irises have exceptionally large, dark-colored flowers, associated with night-sheltering pollination and heat reward by the dark flowers., We quantified phenotypic selection on stem length, floral size and color in two species of iris ( Iris atropurpurea and I. haynei), using an experimental approach. We estimated selection gradients for both flowers open to natural pollination and for flowers receiving supplementary hand pollination, assuming that open-pollinated flowers are affected by all factors that could influence fitness, whereas supplementary pollination removes the possible influence of pollinators., We found evidence for pollinator-mediated selection to increase floral size and stem length in I. atropurpurea, but floral color in this species was not under pollinator-mediated selection. In I. haynei, no pollinator-mediated selection on any of the traits was detected., We conclude that the extreme floral size of I. atropurpurea has probably evolved as a result of pollinator behavior. Lack of such evidence for I. haynei and for the dark floral color in both species suggests that other non-pollinator agents are selecting for these prominent traits, or that phenotypic color variation in these irises is neutral. [ABSTRACT FROM AUTHOR]

Larger and longer lived flowers receive more pollinators, but may also involve increased water maintenance costs under hot, dry environments. Hence, smaller and/or short-lived flowers may buffer such costs. We surveyed floral longevity in three large-flowered MediterraneanCistusspecies. We hypothesize that: (1) inCistus, floral longevity decreases with increasing air temperature and flower size; (2) inC. ladanifer, flower size and longevity increase along an altitudinal gradient; (3) floral longevity is differentially affected by temperature rather than flower size along the gradient; (4) under similar temperature, floral longevity decreases with flower size. For each species, we evaluated the effects of flower size and air temperature on floral longevity. Specifically, floral longevity was surveyed along an altitudinal gradient in the largest flowered speciesCistusladanifer. Floral longevity inCistusspecies lasted < 1 d and was affected by air temperature, independently of flower size. InC. ladanifer, flower size increased along the gradient but floral longevity decreased. Still, floral longevity decreased with increasing air temperature and, to a lesser extent, with flower size. Together, our findings show a triangular relationship among air temperature, flower size and floral longevity with margins for plasticity to accommodate pollinator attraction with the costs of large-flowered Mediterranean plants. [ABSTRACT FROM AUTHOR]

Flower size and number usually evolve under pollinator-mediated selection. However, hot, dry environments can also modulate display, counteracting pollinator attraction. Increased pollen deposition on larger flower displays may not involve higher female fitness. Consequently, stressful conditions may constrain flower size, favouring smaller-sized flowers. The large-flowered, self-incompatible Mediterranean shrub Cistus ladanifer was used to test that: (1) this species suffers pollen limitation; (2) pollinators are spatially-temporally variable and differentially visit plants with more/larger flowers; (3) increased visits enhance reproduction under pollen limitation; (4) stressful conditions reduce female fitness of larger displays; and (5) phenotypic selection on floral display is not just pollinator-mediated. We evaluated pollen limitation, related floral display to pollinator visits and fruit and seed production and estimated phenotypic selection. Flower size was 7.2-10.5 cm and varied spatially-temporally. Visitation rates (total visits/50 min) ranged from 0.26 to 0.43 and increased with flower size. Fruit set averaged 80% and seed number averaged 855, but only fruit set varied between populations and years. Selection towards larger flowers was detected under conditions of pollen limitation. Otherwise, we detected stabilizing selection on flower size and negative selection on flower number. Our results suggest that selection on floral display is not only pollinator-dependent through female fitness in C. ladanifer. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 540-555. [ABSTRACT FROM AUTHOR]

Background Animals fertilize thousands of angiosperm species whose floral-display sizes can significantly influence pollinator behavior and plant reproductive success. Many studies have measured the interactions among pollinator behavior, floral-display size, and plant reproductive success, but few studies have been able to separate the effects of pollinator behavior and post-pollination processes on angiosperm sexual reproduction. In this study, we utilized the highly self-incompatible pollinium-pollination system of Asclepias syriaca (Common Milkweed) to quantify how insect visitors influenced male reproductive success measured as pollen removal, female reproductive success measured as pollen deposition, and self-pollination rate. We also determined how floral-display size impacts both visitor behavior and self-pollination rate. Results Four insect taxonomic orders visited A. syriaca: Coleoptera, Diptera, Hymenoptera, and Lepidoptera. We focused on three groups of visitor taxa within two orders (Hymenoptera and Lepidoptera) with sample sizes large enough for quantitative analysis: Apis mellifera (Western Honey Bee), Bombus spp. (bumble bees) and lepidopterans (butterflies and moths). Qualitatively, lepidopterans had the highest pollinator importance values, but the large variability in the lepidopteran data precluded meaningful interpretation of much of their behavior. The introduced A. mellifera was the most effective and most important diurnal pollinator with regard to both pollen removal and pollen deposition. However, when considering the self-incompatibility of A. syriaca, A. mellifera was not the most important pollinator because of its high self-pollination rate as compared to Bombus spp. Additionally, the rate of self-pollination increased more rapidly with the number of flowers per inflorescence in A. mellifera than in the native Bombus spp. Conclusions Apis mellifera's high rate of self-pollination may have significant negative effects on both male and female reproductive successes in A. syriaca, causing different selection on floraldisplay size than native pollinators. [ABSTRACT FROM AUTHOR]

Large floral displays receive more pollinator visits but involve higher production and maintenance costs. This can result in indirect costs which may negatively affect functions like reproductive output. In this study, we explored the relationship between floral display and indirect costs in two pairs of coflowering sympatric Mediterranean Cistus of contrasting flower size. We hypothesized that: (1) corolla production entails direct costs in dry mass, N and P, (2) corollas entail significant indirect costs in terms of fruit set and seed production, (3) indirect costs increase with floral display, (4) indirect costs are greater in larger-flowered sympatric species, and (5) local climatic conditions influence indirect costs. We compared fruit set and seed production of petal-removed flowers and unmanipulated control flowers and evaluated the influence of mean flower number and mean flower size on relative fruit and seed gain of petal-removed and control flowers. Fruit set and seed production were significantly higher in petal-removed flowers in all the studied species. A positive relationship was found between relative fruit gain and mean individual flower size within species. In one pair of species, fruit gain was higher in the large-flowered species, as was the correlation between fruit gain and mean number of open flowers. In the other pair, the correlation between fruit gain and mean flower size was also higher in the large-flowered species. These results reveal that Mediterranean environments impose significant constraints on floral display, counteracting advantages of large flowers from the pollination point of view with increased indirect costs of such flowers. [ABSTRACT FROM AUTHOR]

Premise of research. In flowers that are specialized, floral traits often differentiate in association with pollinator shifts. However, in species of generalized pollination, the role of pollinators in local population differentiation of floral traits is difficult to test. A crucial line of evidence is to contrast the direction and strength of pollinator selection among populations. Methodology. Variation in floral traits and pollinator assemblages was investigated in two populations of a generalist-pollinated herb, Trollius ranunculoides (Ranunculaceae), to detect the potential role of pollinators in floral differentiation. Pivotal results. The alpine population was visited more by flies, and the subalpine population was visited more by bees. Flower manipulations suggested that large flowers were more attractive to both bees and flies, while more petals were favored only by nectar-feeding flies. Compared to the subalpine population, plants in the alpine population had more petals per flower but smaller flowers where flies were predominant pollinators. Flower differences appeared to be genetically maintained in a common garden. Phenotypic selection analysis indicated stronger selection on petal number in the alpine population and oppositely more intense selection on flower diameter in the subalpine population, in accordance with the preferences of local pollinator assemblages. Conclusions. Heritability in floral traits, differences in local pollinator assemblages, and corresponding differences in phenotypic selection are together consistent with the view of pollinator-mediated selection on floral differentiation in T. ranunculoides. [ABSTRACT FROM AUTHOR]

Floral size dimorphism, pollination, and genetic variation of Alpinia nieuwenhuizii (Zingiberaceae), a flexitylous ginger, were studied. This study revealed that floral size differed among habitats (i.e., roadsides/riversides vs. forest floors). The effective pollinators of small-flowered populations of the species on a forest floor were different from those of large-flowered populations along roadsides/riversides. Using inter-simple sequence repeat (ISSR) PCR, considerable genetic differentiation was detected between small- and large-flowered populations. These results indicate that reproductive isolation in A. nieuwenhuizii owing to the differentiation of pollen vectors between two floral size morphs may lead to genetic differentiation between the two morphs. [ABSTRACT FROM AUTHOR]

Abstract: Plants in small populations may receive fewer visits, smaller pollen loads or pollen of poorer quality and suffer from reduced reproductive success compared to plants in larger populations. Consequently, pollen limitation of plants in small populations has been suggested to result in the evolution of reduced reliance on pollinators or the enhancement of traits that attract pollinators. The main aim of this study was to experimentally quantify the strength of pollinator-mediated selection on floral display size and flowering phenology in populations of varying size, using the self-incompatible, perennial herb Phyteuma spicatum as study species. We conducted supplementary hand pollinations in six populations (ranging in size between ca. 20–3000 flowering individuals) over two consecutive years and assessed selection gradients (i.e., trait–fitness relationships) in open- and hand-pollinated plants. Our results show that some populations are pollen limited in some years, but, contrary to our expectation, the degree of pollen limitation was not significantly related to population size. We found phenotypic selection for increased inflorescence size (in most populations and in both years), but we obtained no or no strong evidence that selection was pollinator-mediated or that the strength of selection was related to population size. This may have been the result of low statistical power, an inherent problem of studies examining effects of population size that require the inclusion of populations with only few individuals. In addition, given that selection appeared to be spatially and temporally variable, abiotic or biotic factors other than pollinators may have contributed to selection on inflorescence size. [Copyright &y& Elsevier]

Abstract: Fluctuating conditions throughout the year and changes in floral display may promote shifts in pollinator activity and predator pressure, influencing female reproductive output, especially for extended flowering species under seasonal climates. In this regard, flowering and fruit production were tracked in 2 different years in 2 populations of Ononis tridentata in Central Spain. Total fruit production was estimated, and fruits were harvested to obtain primary fruit investment, pollination success, predation incidence, seed production, seed weight and germination rates. Ononis tridentata combined spring mass flowering with a steady long flowering period across the summer and fall. The fewer flowers that are produced in fall were successfully pollinated, and produced fruits that were subject to minimal predation pressure relative to spring fruits. Moreover, fall fruits contained a higher number of heavier seeds and showed higher germination rates than those of spring seeds. Fall reproductive output represent 10% of annual viable seeds and thus may act as an important complement to the main spring reproductive investment. Extended flowering could be interpreted as a “bet-hedging strategy” for enduring Mediterranean unpredictable and changing environmental conditions. [Copyright &y& Elsevier]

Floral traits used to attract and reward pollinators should become vestigial during the evolution of self-fertilization from outcrossing due to the reduced fitness benefits of investing in pollinator attraction. However, few traits have been studied, and covariation with outcrossing is rarely evaluated using genetic estimates of the mating system. We examined covariation between petal size and color and genetic estimates of outcrossing among populations of the yellow-flowered, bee-pollinated Camissoniopsis cheiranthifolia that exhibit striking variation in the mating system. We measured brightness, chroma, and hue, using a spectrophotometer, and petal size, using image analysis for flowers sampled in the field from 25 populations across the species' geographic range. Petals were brighter with higher chroma and lower hue (less red) in large-flowered populations than in small-flowered populations. Petal size, brightness, and chroma also correlated positively with the proportion of seeds outcrossed estimated from marker-gene analysis for a subsample of 11 populations. Parallel changes in petal size and color in selfing populations from two climatically different regions plus the maintenance of differences in flower size and color between large- and small-flowered populations when grown in a common glasshouse environment suggests genetic differentiation in these traits associated with mating system evolution. [ABSTRACT FROM AUTHOR]

Although pollinators are thought to select on flower colour, few studies have experimentally decoupled effects of colour from correlated traits on pollinator visitation and pollen transfer. We combined selection analysis and phenotypic manipulations to measure the effect of petal colour on visitation and pollen export at two spatial scales in Wahlenbergia albomarginata. This species is representative of many New Zealand alpine herbs that have secondarily evolved white or pale flowers. The major pollinators, solitary bees, exerted phenotypic selection on flower size but not colour, quantified by bee vision. When presented with manipulated flowers, bees visited flowers painted blue to resemble a congener over white flowers in large, but not small, experimental arrays. Pollen export was higher for blue flowers in large arrays. Pollinator preference does not explain the pale colouration of W. albomarginata, as commonly hypothesized. Absence of bright blue could be driven instead by indirect selection of correlated characters. [ABSTRACT FROM AUTHOR]

Background and Aims Dimorphism among floral traits can evolve through variation in selection intensity between female and male performance, especially when sex functions are separated between flowers on a plant (monoecy), or between individuals (dioecy). In animal-pollinated species, male floral traits are predicted to be larger because competition for pollinators should favour larger displays. Floral dimorphism may be greater in dioecious than monoecious populations because of trade-offs between female and male function and opportunities for selfing in hermaphrodites. Methods These predictions were tested by surveying flower size, total flowers per inflorescence and daily display size in the insect-pollinated Sagittaria latifolia (Alismataceae). This species is useful for comparative analysis because populations are mostly either monoecious or dioecious. We examined floral dimorphism in 13 monoecious and 16 dioecious populations in eastern North America. Key Results Male flowers were significantly larger than female flowers in monoecious and dioecious populations, but there was no evidence for greater flower size dimorphism in dioecious populations despite their larger flower sizes overall. Although inflorescences in both dioecious and monoecious populations produced more male flowers, daily floral displays were significantly larger for female than male function due to more synchronous female flower opening. Daily floral display dimorphism was significantly greater in dioecious populations, due to greater female daily floral displays. There was a positive relationship between mean flower size and total flowers per inflorescence for both sexes in dioecious populations, but no relationship for either sex function in monoecious populations. Flower size dimorphism was positively correlated with the frequencies of females in dioecious populations. Conclusions The increased size and number of male flowers and protracted male floral displays in S. latifolia are probably shaped by sexual selection for more effective pollen dispersal. [ABSTRACT FROM PUBLISHER]

The rate of pollen exchange within and among flowers may depend on pollinator attraction traits such as floral display size and flowering plant density. Variations in these traits may influence pollinator movements, pollen receipt, and seed number. To assess how floral display size and flowering plant density affect parameters of pollinator visitation rate, pollen receipt per flower, seed number per fruit and the between-plant pollinator movements, we studied the self-incompatible plant, Nierembergia linariifolia. Per-flower pollinator visitation rate and bout length increased linearly with increasing floral display size. Pollen receipt per flower increased linearly with increasing flowering plant density. For seed number per fruit, a polynomial model describing an increased seed number per fruit at low density and a decreased seed number per fruit at high density provided a significant fit. Per-flower pollinator visitation rate was not associated with pollen receipt per flower and seed number per fruit. Bees visited plants located near to the center of the population more frequently than plants located at the periphery. Increases in both floral display size and flowering plant density led to an increased chance of a plant being chosen as the center of the pollinator foraging area. These results suggest that even though large floral displays and high flowering plant density are traits that attract more pollinators, they may also reduce potential mate diversity by restricting pollen movement to conspecific mates that are closely located. [ABSTRACT FROM AUTHOR]