Simulation analyses of the evolution of intra-inflorescence flowering patterns assuming selection on anthesis interval among individual flowers.

What conditions select flowering patterns within inflorescences, or variation in the anthesis interval within inflorescences among plants? Under what conditions are gradual blooming and simultaneous blooming, both traits related to floral display size, advantageous? We constructed a simulation model in which the opening times and longevities of individual flowers within inflorescences, the sizes of attractive structures of individual flowers, and the numbers of ovules and pollen grains produced by individual flowers evolve. Individual plants in the population compete for pollinators, and plants are selected by pollinators according to their floral display sizes and amounts of resources allocated to attractive structures. We found that, if the proportion of pollen on a pollinator deposited on a stigma was low, gradual blooming did not evolve even if inbreeding depression was greater than 0.5. This is because the amount of outcross-pollen on pollinators decreased at a low rate during flower visits within a single inflorescence, and the selfing rate was suppressed to a low level even if the floral display size was large. On the other hand, if the proportion of pollen deposition was high, gradual blooming evolved even if inbreeding depression was smaller than 0.5. This may be because gradual blooming can enhance pollen delivery to other plants by reducing the loss of self-pollen by geitonogamy. On the other hand, allocation ratios among floral organs (female and male organs and attractive structures) were independent of the degree of simultaneous and gradual blooming within inflorescences. We concluded that the evolution of gradual blooming is more strongly affected by the proportion of pollen on a pollinator deposited on a stigma than by inbreeding depression.

Variations of floral temperature in changing weather conditions.

Floral temperature is a flower characteristic that has the potential to impact the fitness of flowering plants and their pollinators. Likewise, the presence of floral temperature patterns, areas of contrasting temperature across the flower, can have similar impacts on the fitness of both mutualists. It is currently poorly understood how floral temperature changes under the influence of different weather conditions, and how floral traits may moderate these changes. The way that floral temperature changes with weather conditions will impact how stable floral temperatures are over time and their utility to plants and pollinators. The stability of floral temperature cues is likely to facilitate effective plant-pollinator interactions and play a role in the plant's reproductive success. We use thermal imaging to monitor how floral temperatures and temperature patterns of four plant species (Cistus 'snow fire' and 'snow white', Coreopsis verticillata and Geranium psilostemon) change with several weather variables (illumination, temperature; windspeed; cloud cover; humidity and pressure) during times that pollinators are active. All weather variables influenced floral temperature in one or more species. The directionality of these relationships was similar across species. In all species, light conditions (illumination) had the greatest influence on floral temperatures overall. Floral temperature and the extent to which flowers showed contrasting temperature patterns were influenced predominantly by light conditions. However, several weather variables had additional, lesser, influences. Furthermore, differences in floral traits, pigmentation and structure, likely resulted in differences in temperature responses to given conditions between species and different parts of the same flower. However, floral temperatures and contrasting temperature patterns that are sufficiently elevated for detection by pollinators were maintained across most conditions if flowers received moderate illumination. This suggests the presence of elevated floral temperature and contrasting temperature patterns are fairly constant and may have potential to influence plant-pollinator interactions across weather conditions.

Among-individual variation in flowering phenology affects flowering synchrony and mating opportunity.

PREMISE: The timing and pattern of a plant's flowering can have important consequences for reproductive success. Variation in flowering phenology may influence the number of prospective mates, the risk of mating with lower quality individuals, and the likelihood of self-pollination. Here we use a common garden experiment to explore within- and among-population variation in phenology. Our work provides new insights into how flowering phenology shapes mating opportunity and flowering synchrony in a self-compatible perennial. METHODS: To quantify variation in flowering phenology we raised progeny from nine populations of Mimulus ringens in a common garden. For each individual, we measured phenological traits including age at flowering onset, daily floral display size, total flower number, and flowering synchrony with other members of the population, and related these traits to mating opportunity. We also tested how individual flowering schedules influence the magnitude of synchrony. RESULTS: Flowering phenology and synchrony varied substantially within and among populations. From day to day, plants often oscillated between large and small daily floral displays. Additionally, flowering schedules of individual plants strongly influenced flowering synchrony and, along with the number of flowering days, markedly affected plants' mating opportunity. CONCLUSIONS: Phenological traits such as flowering synchrony can affect the quantity of mating opportunities and may be important targets of natural selection. Our results highlight the need for studies that quantify flowering patterns of individuals as well as populations.

In Which Way Do the Flower Properties of the Specialist Orchid Goodyera repens Meet the Requirements of Its Generalist Pollinators?

This article is the next part of a series of studies documenting the influence of flower traits on the reproductive success (RS) of orchids. Knowledge of factors influencing RS helps to understand the mechanisms and processes crucial for shaping plant-pollinator interactions. The aim of the present study was to determine the role of flower structure and nectar composition in shaping the RS of the specialist orchid Goodyea repens, which is pollinated by generalist bumblebees. We found a high level of pollinaria removal (PR) and female reproductive success (fruiting, FRS) as well as a high level of variation between populations, although in certain populations pollination efficiency was low. Floral display traits, mainly inflorescence length, influenced FRS in certain populations. Among the flower traits, only the height of flowers was correlated with FRS in one population, suggesting that the flower structure of this orchid is well adapted to pollination by bumblebees. The nectar of G. repens is diluted and dominated by hexoses. Sugars were less important in shaping RS than amino acids. At the species level, twenty proteogenic and six non-proteogenic AAs were noted, along with their differentiated amounts and participation in particular populations. We found that distinct AAs or their groups mainly shaped PR, especially when correlations were considered at the species level. Our results suggest that both the individual nectar components and the ratios between them have an impact on G. repens RS. Because different nectar components influence the RS parameters in different ways (i.e., negatively or positively), we suggest that different Bombus species play the role of main pollinators in distinct populations.

Does Reproductive Success in Natural and Anthropogenic Populations of Generalist Epipactis helleborine Depend on Flower Morphology and Nectar Composition?

The purpose of our study was to determine the role of flower structure and nectar composition in shaping the reproductive success (RS) of the generalist orchid Epipactis helleborine in natural and anthropogenic populations. We supposed that the distinct character of two groups of habitats creates different conditions for plant-pollinator relationships, thus influencing reproductive success in E. helleborine populations. Both pollinaria removal (PR) and fruiting (FRS) were differentiated between the populations. On average, FRS was almost two times higher in the anthropogenic than in the natural populations. The difference between the two population groups in PR was smaller but still statistically significant. RS parameters were correlated with some floral display and flower traits. Floral display influenced RS only in three anthropogenic populations. Flower traits had a weak influence on RS (10 of the 192 cases analyzed). The more important trait in shaping RS was nectar chemistry. The nectar of E. helleborine is relatively diluted with a lower sugar concentration in the anthropogenic than in the natural populations. In the natural populations, domination of sucrose over hexoses was found, while in the anthropogenic populations, hexoses were more abundant and the participation of sugars was balanced. In some populations, sugars influenced RS. In E. helleborine nectar, 20 proteogenic and 7 non-proteogenic amino acids (AAs) were found with a clear domination of glutamic acid. We noted relationships between some AAs and RS, but distinct AAs shaped RS in different populations, and their impact was independent of their participation. Our results indicate that the flower structure and nectar composition of E. helleborine reflect its generalistic character and meet the requirements of a wide range of pollinators. Simultaneously, the differentiation of flower traits suggests a variation in pollinator assemblages in particular populations. Knowledge about the factors influencing RS in distinct habitats helps to understand the evolutionary potential of species and to understand mechanisms and processes crucial for shaping interactions between plants and pollinators.

Effect of grazing disturbance on floral display, pollen limitation and plant pollination efficiency in the desert steppe.

BACKGROUND: Grazing disturbance usually affects floral display and pollination efficiency in the desert steppe, which may cause pollen limitation in insect-pollinated plants. Effective pollination is essential for the reproductive success of insect-pollinated plants and insufficient pollen transfer may result in pollen limitation. Caragana microphylla Lam is an arid region shrub with ecological importance. Few studies have been conducted on how grazing disturbance influences pollen limitation and pollination efficiency of C. microphylla. Here, we quantify the effect of different grazing intensities on floral display, pollinator visitation frequency and seed production in the Urat desert steppe. RESULTS: In C. microphylla, supplemental hand pollination increased the seed set, and pollen limitation was the predominant limiting factor. As the heavy grazing significantly reduced the seed set in plants that underwent open-pollination, but there was no significant difference in the seed set between plants in the control plots and plants in the moderate grazing plots. Furthermore, there was a higher pollinator visitation frequency in plants in the control plots than in plants in the heavy grazing plots. CONCLUSIONS: We found that pollinator visitation frequency was significantly associated with the number of open flowers. Our findings also demonstrated that seed production is associated with pollinator visitation frequency, as indicated by increased seed production in flowers with higher pollinator visitation frequency. Therefore, this study provides insight into the effect of different grazing intensities on floral display that are important for influencing pollinator visitation frequency and pollination efficiency in desert steppes.

Disruptive selection via pollinators and seed predators on the height of flowers on a wind-dispersed alpine herb.

PREMISE: Floral stalk height is known to affect seed dispersal of wind-dispersed grassland species, but it may also affect the attractiveness of flowers and fruits of animal-pollinated and animal-dispersed plants. Stalk height may thus be responsive to selection via interactions with both mutualist pollinators and seed dispersers, but also antagonist florivores and seed predators. In this study, we aimed to determine the effect of pollinators and seed predators on selection on floral stalk height in the insect-pollinated and wind-dispersed, alpine, andromonoecious herb Pulsatilla alpina, whose flowers also vary in their sex allocation and thus in the resources available to both mutualists and antagonists. METHODS: We measured the resource status of individuals in terms of their size and the height of the vegetation surrounding plants of P. alpina at 11 sites. In one population, we recorded floral stalk height over an entire growing season and investigated its association with floral morphology and floral sex allocation (pistil and stamen number) and used leaf-removal manipulations to assess the effect of herbivory on floral stalk height. Finally, in four populations, we quantified phenotypic selection on floral stalk height in four female components of reproductive success before seed dispersal. RESULTS: Stalk height was positively associated with female allocation of the respective flower, the resource status of the individual, and the height of the surrounding vegetation, and negatively affected by leaf removal. Our results point to disruptive selection on stalk height in terms of both selection differentials and selection gradients for fertilization, seed predation, and seed maturation rates and to positive selection on stalk height in terms of a selection differential for mature seed number. CONCLUSIONS: Stalk height of P. alpina is a costly trait that affects female reproductive success via interactions with both mutualists and antagonists. We discuss the interplay between the resource status and selection imposed on female reproductive success and its likely role in the evolution of sex-allocation strategies, especially andromonoecy.

Pollen transfer and patterns of reproductive success in pure and mixed populations of nectariferous Platanthera bifolia and P. chlorantha (Orchidaceae).

Plant species evolution is driven by many factors that have different roles in space and time. Using different field and laboratory methods, we studied reproductive patterns and their determinants in pure and mixed P. bifolia and P. chlorantha populations in different habitats. We also considered the probability of hybridisation between these two species and the role of intra-population processes in maintaining species integrity. Generally, we found a high level of reproductive success in both Platantherans. In both species, male (MRS) and female (FRS) reproductive success depended on floral display, and male reproductive success additionally on population structure. The flower traits were only weakly related to reproductive success. Moths' assemblages varied spatially and temporally, and their diversity and numbers were correlated with MRS in the year, when their abundance was markedly lower. Analysis of patterns of pollen transfer showed that pollen was transported up to 25 m (average 8.2 ± 4.83 m) and showed gene exchange between these two Platanthera species. The germination level of both species was significantly lower than seed viability, although P. bifolia seed germinated with higher frequency than P. chlorantha seeds. We noted differences in viability and germination of seeds developed as an effect of experimental interspecies crossings and those developed from natural pollination. The presence of intermediate ecotypes together with observations of spontaneous interspecies crosses in the field and viability of seeds produced in interspecies crossing suggest that both pre- and postzygotic reproductive barriers are not complete and do not prevent hybrid production.

Plasticity in floral longevity and sex-phase duration of Lobelia siphilitica in response to simulated pollinator declines.

PREMISE: Pollinator declines can reduce the quantity and quality of pollination services, resulting in less pollen deposited on flowers and lower seed production by plants. In response to these reductions, plant species that cannot autonomously self-pollinate and thus are dependent on pollinators to set seed could plastically adjust their floral traits. Such plasticity could increase the opportunity for outcross pollination directly, as well as indirectly by affecting inflorescence traits. METHODS: To test whether plants can respond to pollinator declines by plastically adjusting their floral traits, we simulated declines by experimentally reducing pollinator access to Lobelia siphilitica plants and measuring two traits of early- and late-season flowers: (1) floral longevity; and (2) sex-phase duration. To test whether plasticity in these floral traits affected inflorescence traits, we measured daily display size and phenotypic gender. RESULTS: We found that experimentally reducing pollination did not affect female-phase duration, but did extend the male-phase duration of early-season flowers by 13% and the longevity of late-season flowers by 12.8%. However, plants with an extended male phase did not have a more male-biased phenotypic gender, and plants with an extended floral longevity did not have a larger daily display. CONCLUSIONS: Our results suggest that plants can respond to pollinator declines by plastically adjusting both the longevity and sex-phase duration of their flowers. If this plasticity increases the opportunity for outcross pollination, then it could be one mechanism by which pollinator-dependent plant species maintain seed production as pollinators decline.

Nectar Chemistry or Flower Morphology-What Is More Important for the Reproductive Success of Generalist Orchid Epipactis palustris in Natural and Anthropogenic Populations?

The aim of this study was to determine the level of reproductive success (RS) in natural and anthropogenic populations of generalist orchid Epipactis palustris and its dependence on flower structure and nectar composition, i.e., amino acids and sugars. We found that both pollinaria removal and female reproductive success were high and similar in all populations, despite differences in flower traits and nectar chemistry. Flower structures were weakly correlated with parameters of RS. Nectar traits were more important in shaping RS; although, we noted differentiated selection on nectar components in distinct populations. Individuals in natural populations produced nectar with a larger amount of sugars and amino acids. The sucrose to (fructose and glucose) ratio in natural populations was close to 1, while in anthropogenic ones, a clear domination of fructose and glucose was noted. Our results indicate that the flower traits and nectar composition of E. palustris reflect its generalist character and meet the requirements of a wide range of pollinators, differing according to body sizes, mouth apparatus, and dietary needs. Simultaneously, differentiation of nectar chemistry suggests a variation of pollinator assemblages in particular populations or domination of their some groups. To our knowledge, a comparison of nectar chemistry between natural and anthropogenic populations of orchids is reported for the first time in this paper.

Self-Fertilization, Inbreeding, and Yield in Alfalfa Seed Production.

Selfing (self-pollination) is the ultimate form of inbreeding, or mating among close relatives. Selfing can create yield loss when inbreeding depression, defined as a lower survival and reproduction of inbred relative to outbred progeny, is present. To determine the impact of selfing in alfalfa (Medicago sativa L.), we quantified the selfing rate of 32 alfalfa seed production fields located in three regions, namely, the Pacific Northwest (PNW), the Central Valley of California (CEV), and the Imperial Valley of California (IMP). Selfing rates (the proportion of selfed seeds) varied between 5.3 and 30% with an average of 12.2% over the 32 seed production fields. In both the parents and their progeny, we observed an excess of heterozygotes relative to Hardy-Weinberg expectations. We detected notable levels of inbreeding in parents (0.231 ± 0.007 parental inbreeding coefficient) and progeny (0.229 ± 0.005). There were a 15% decrease in the number of seeds per stem (seed set) and a 13% decline in the number of seeds per pod in selfed relative to outcrossed stems, but negligible inbreeding depression for pods per raceme and seed weight. The number of racemes on selfed stems increased significantly in fields with greater selfing rates, supporting the presence of geitonogamous or among flower selfing. Despite the significant level of inbreeding depression, seed set did not decrease in fields with higher selfing rates, where the greater number of racemes on the selfed stems increased the seed set. The effects of the field selfing rate on the seed yield metrics were mostly indirect with direct effects of the number of racemes per stem. Available data indicate that the majority of selfing in alfalfa is pollinator-mediated, and thus, eliminating selfing in alfalfa seed production would require the selection of self-incompatible varieties, which, by eliminating inbreeding depression, would provide a 15% potential increase in seed yield and an increase in future hay yield.

Bumblebees can detect floral humidity.

Floral humidity, a region of elevated humidity in the headspace of the flower, occurs in many plant species and may add to their multimodal floral displays. So far, the ability to detect and respond to floral humidity cues has been only established for hawkmoths when they locate and extract nectar while hovering in front of some moth-pollinated flowers. To test whether floral humidity can be used by other more widespread generalist pollinators, we designed artificial flowers that presented biologically relevant levels of humidity similar to those shown by flowering plants. Bumblebees showed a spontaneous preference for flowers that produced higher floral humidity. Furthermore, learning experiments showed that bumblebees are able to use differences in floral humidity to distinguish between rewarding and non-rewarding flowers. Our results indicate that bumblebees are sensitive to different levels of floral humidity. In this way floral humidity can add to the information provided by flowers and could impact pollinator behaviour more significantly than previously thought.

Selfing rates vary with floral display, pollinator visitation and plant density in natural populations of Mimulus ringens.

Variation in selfing rates within and among populations of hermaphroditic flowering plants can strongly influence the evolution of reproductive strategies and the genetic structure of populations. This intraspecific variation in mating patterns may reflect both genetic and ecological factors, but the relative importance of these factors remains poorly understood. Here, we explore how selfing in 13 natural populations of the perennial wildflower Mimulus ringens is influenced by (a) pollinator visitation, an ecological factor, and (b) floral display, a trait with a genetic component that also responds to environmental variation. We also explore whether genetically based floral traits, including herkogamy, affect selfing. We found substantial variation among populations in selfing rate (0.13-0.55). Selfing increased strongly and significantly with floral display, among as well as within populations. Selfing also increased at sites with lower pollinator visitation and low plant density. However, selfing was not correlated with floral morphology. Overall, these results suggest that pollinator visitation and floral display, two factors that interact to affect geitonogamous pollinator movements, can influence the selfing rate. This study identifies mechanisms that may play a role in maintaining selfing rate variation among populations.

Asymmetry in fitness-related traits of later-generation hybrids between two invasive species.

PREMISE: The importance of hybridization to invasion has been frequently discussed, with most studies focusing on the comparison of fitness-related traits between F1 hybrids and their parents and the consequences of such fitness differences. However, relatively little attention has been given to late-generation hybrids. Different fitness landscapes could emerge in later generations after hybrids cross with each other or backcross with their parents, which may play an important role in plant invasion and subsequent speciation. METHODS: In this study, artificial crosses were conducted to generate multiple generations, including F1, F2, and backcrosses between two invasive species: Cakile edentula (self-compatible) and C. maritima (self-incompatible). Putative hybrids were also collected in the sympatric zone and compared with their co-occurring parents for phenotypic and genetic differences. RESULTS: Genetic data provided evidence of hybridization happening in the wild, and phenotypic comparisons showed that natural hybrids had intermediate traits between the two species but showed more similarity to C. maritima than to C. edentula. The asymmetry was further identified in artificial generations for several phenotypic characters. Furthermore, backcrosses exhibited different patterns of variation, with backcrosses to C. maritima having higher reproductive output than their counterparts. CONCLUSIONS: Our results suggest that hybridization and introgression (backcrossing) in Cakile species is asymmetric and most likely to favor the proliferation of C. maritima genes in the mixed population and thus help its establishment, a finding that could not be predicted by characterizing F1 hybrids.

The effects of pollination, herbivory and autonomous selfing on the maintenance of flower colour variation in Silenelittorea.

Intraspecific flower colour variation has been generally proposed to evolve as a result of selection driven by biotic or abiotic agents. In a polymorphic population of Silene littorea with pink- and white-flowered plants, we studied pollinators, analysed flower colour perception and tested for differences in pollinator visitation. We also experimentally analysed pollinator limitation in fruit and seed set, and the degree of autonomous selfing. The incidence of florivory and leaf herbivory was compared over 3-4 years. Silene littorea is mainly pollinated by bees and butterflies. Pollinators preferred pink flowers, which did not show pollinator limitation. On the contrary, white flowers showed pollinator limitation in fruit set. White-flowered plants had less floral display and higher levels of florivory than pink plants. Flower colour morphs of S. littorea can reproduce in the absence of pollinators by autonomous selfing, setting 20% and 12% of fruit and seeds in the pink morph and 27% and 20% in the white morph, respectively. Fruit set of white flowers produced by autonomous selfing did not differ from open-pollinated flowers. In conclusion, S. littorea is pollinated by insects of different orders that more frequently visit pink flowers, which is reflected in pollinator limitation of fruit set in white flowers. Moreover, this species has a mixed mating system in which both colour morphs can reproduce in the absence of pollinators by autonomous selfing, although white flowers mainly produce fruits by autogamy. We suggest that reproductive assurance by autonomous selfing helps to maintain flower colour polymorphism in this population.

Below-ground competition alters attractiveness of an insect-pollinated plant to pollinators.

Competitive interactions between plants can affect patterns of allocation to reproductive structures through modulation of resource availability. As floral traits involved in plant attractiveness to pollinators can be sensitive to these resources, competition with any neighbouring species may influence the attractiveness of insect-pollinated plants. While pollination research has primarily focused on above-ground interactions, this study aims at investigating if the presence of a competitor plant can modulate neighbouring insect-pollinated plant attractiveness to pollinators and resulting fecundity, especially through below-ground competitive interactions for soil resources. We set up a plot experiment in which we grew an insect-pollinated plant, Sinapis alba (Brassicaceae), in a mixture dominated by a wind-pollinated plant, Holcus lanatus (Poaceae). Individuals of S. alba were either subjected to or isolated from (with buried tubes in the soil) below-ground competition. Across the flowering season, floral traits involved in attractiveness of S. alba and pollinator visitation were followed at the plot and plant level to investigate different scales of attractiveness. At the end of the experiment, seeds were harvested to assess plant fecundity. Competition had a significant negative effect on plot and plant floral display size as well as flower size while nectar traits were not affected. When plants of S. alba were in competition, the time to first visit was altered: the proportion of plots that received a visit was smaller for a given time; in other words, it took more time for a given proportion of plots to be visited and some plots were even never visited. Moreover, pollinators made fewer visits per plots. The proportion of viable seeds produced by S. alba in competition was lower and probably linked to the competition itself rather than changes in pollinator visitation. This study suggests that competitive interactions between plants can modulate pollination interactions even when competing plant species are not insect-pollinated.

The Influence of Arbuscular Mycorrhizal Fungi on Plant Reproduction.

Arbuscular mycorrhizal (AM) fungi can influence all components of plant reproduction including pollen delivery, pollen germination, pollen tube growth, fertilization, and seed germination. AM fungi associate with plant roots, uptake nutrients, and prime plants for faster defense responses. Our literature review first identified four testable hypotheses describing how AM fungi could alter pollen delivery: (1) We hypothesize AM fungi promote floral display size. The influence of AM fungi on flower size and number is supported by literature, however there are no studies on floral color. (2) We hypothesize AM fungi promote pollen and nectar quality and quantity, and, as reported before, AM fungi promote male fitness over female fitness. (3) We hypothesize AM fungi promote both earlier and longer flowering times, but we found no consistent trend in the data for earlier or later or longer flowering times. (4) We hypothesize AM fungi alter floral secondary chemistry and VOCs, and find there is clear evidence for the alteration of floral chemistry but little data on VOCs. Second, we focus on how AM fungi could alter pollen germination, pollen tube growth, and fertilization, and present three testable hypotheses. We found evidence that AM fungi influence pollen germination and pollen tube growth, production of seeds, and seed germination. However, while most of these influences are positive they are not conclusive, because studies have been conducted in small numbers of systems and groups. Therefore, we conclude that the majority of research to date may not be measuring the influence of AM fungi on the most important components of plant reproduction: pollen germination, pollen tube growth, fertilization, and seed germination.

Floral Trait Evolution of Angiosperms on Pacific Islands.

Animals frequently evolve unique suites of traits on islands, but whether plants evolve comparable island syndromes remains unresolved. Here, we test the prediction on the basis of natural history observations that insect-pollinated plants evolve smaller flowers on islands than on mainland communities. We examined 556 plant species representing 136 phylogenetically independent contrasts between island and mainland sister taxa. We focused on endemic taxa originating from the Americas associated with seven tropical and subtropical islands of the Pacific Ocean. Contrary to conventional wisdom, flowers were not on average smaller on islands than on the mainland. On specific archipelagos (the Galápagos Islands and Revillagigedo Islands), however, island taxa did evolve smaller flowers. Divergence in flower size between island and mainland taxa also varied among taxonomic families, such that some plant families evolved smaller flowers on islands, other families evolved larger flowers on islands, and some families exhibited no divergence in flower size between island and mainland taxa. Overall, our results show that there is no general island syndrome for flower size, but instead that the evolution of floral morphology is complex and context dependent, depending on variation among islands and plant families. Our results also suggest that if island floras are typically dominated by small flowered species, as suggested by natural history observations, then ecological filtering is a more likely explanation of this pattern than evolutionary divergence postcolonization. We propose future studies that could disentangle the relative roles of ecological filtering and evolution in the distribution of floral traits on islands.