A bee’s eye view of remarkable floral colour patterns in the south-west Australian biodiversity hotspot revealed by false colour photography

Background and AimsColour pattern is a key cue of bee attraction selectively driving the appeal of pollinators. It comprises the main colour of the flower with extra fine patterns, indicating a reward focal point such as nectar, nectaries, pollen, stamens and floral guides. Such advertising of floral traits guides visitation by the insects, ensuring precision in pollen gathering and deposition. The study, focused in the Southwest Australian Floristic Region, aimed to spot bee colour patterns that are usual and unusual, missing, accomplished by mimicry of pollen and anthers, and overlapping between mimic-model species in floral mimicry cases.MethodsFloral colour patterns were examined by false colour photography in 55 flower species of multiple highly diverse natural plant communities in south-west Australia. False colour photography is a method to transform a UV photograph and a colour photograph into a false colour photograph based on the trichromatic vision of bees. This method is particularly effective for rapid screening of large numbers of flowers for the presence of fine-scale bee-sensitive structures and surface roughness that are not detectable using standard spectrophotometry.Key ResultsBee- and bird-pollinated flowers showed the expected but also some remarkable and unusual previously undetected floral colour pattern syndromes. Typical colour patterns include cases of pollen and flower mimicry and UV-absorbing targets. Among the atypical floral colour patterns are unusual white and UV-reflecting flowers of bee-pollinated plants, bicoloured floral guides, consistently occurring in Fabaceae spp., and flowers displaying a selective attractiveness to birds only. In the orchid genera (Diuris and Thelymitra) that employ floral mimicry of model species, we revealed a surprising mimicry phenomenon of anthers mimicked in turn by model species.ConclusionThe study demonstrates the applicability of ‘bee view’ colour imaging for deciphering pollinator cues in a biodiverse flora with potential to be applied to other eco regions. The technique provides an exciting opportunity for indexing floral traits on a biome scale to establish pollination drivers of ecological and evolutionary relevance.     

Pollinator visits to floral colour phases of Fuchsia excorticata

Abstract

Fuchsia excorticata is a gynodioecious tree (endemic to New Zealand) which is pollinated by honeyeater birds. Red, tubular flowers are common among bird-pollinated plants, and the tubular flowers of F. excorticata change colour from green to red. The purpose of the present study was to describe the timing of the colour change, dropping of the floral tube, and nectar production of F. excorticata and to determine how bellbirds (Anthornis melanura) and two introduced species of nectar robbers (Zosterops lateralis and Bombus sp.) respond to the different colour phases.

Floral tubes fell off about 11 days after anthesis in both sexes, with colour change occurring on about Day-4 for female trees and on about Day-5 for hermaphrodite trees. Green-phase hermaphrodite flowers produced significantly more nectar/day than did green-phase female flowers, while red-phase flowers did not produce nectar in either sex. All three floral visitors studied preferentially visited green-phase flowers and virtually ignored the nectarless flowers in the red phase. These results contrast with the general association between red, bird-pollinated flowers and the presence of a nectar reward. We suggest that the non-migratory habit of the New Zealand honeyeaters and the lack of native insect visitors to this species may account for this anomalous green-to-red colour change.     

Short-term effects of burn season on flowering phenology of savanna plants

We examined the effect of season of burn on flowering phenology of groundlayer species, in the year following burns, in a mesic-sand Midwestern oak savanna. Burn treatments were fall, early-season, growing-season, late-season, and 1 or 5 years after a prior early-season wildfire. For these treatments, we compared the number of flowering stems and of flowers for species overall, for the 20 most prolifically flowering species, as well as for species grouped by flowering phenoperiods, and by growth form. Growing-season burn had a significant negative effect on number of flowering stems and total number of flowers. This effect occurred when either the burn occurred during the flowering season or during the season prior to the flowering phenoperiod. Tradescantia ohiensis showed expedited flowering and Phlox pilosa showed delayed flowering in response to early-season burning. Flowering of early shrubs was reduced by the previous fall and early-spring fires, while flowering of mid-season blooming shrubs was reduced by the early- and growing-season burns. Vaccinium and Gaylussacia, early-flowering shrubs, produced fewer flowers 1 year after than 5 years after an early-season burn. Arabis lyrata showed reduced flowering from the early-season burn. We also found four instances where the early-spring burn effect on flowering was more severe than the fall burn effect, suggesting that many frequent early-season burns may be deleterious to flowering and reproduction of some species. Burns occurring too frequently in the same season could negatively affect future flowering and reproduction of these plant species.     

VARIATION IN THE DEGREE OF CLEISTOGAMY WITHIN AND AMONG SPECIES OF THE GRASS DANTHONIA

North American species of the grass Danthonia bear distinct chasmogamous and cleistogamous flowers. Four taxa occurring in North Carolina (D. compressa, D. epilis, D. sericea, and D. spicata) were examined for variation in the degree of cleistogamy. The flowering culms bear a single terminal panicle consisting of 4 to 12 spikelets of chasmogamous flowers. In each leaf axil, at the nodes of the flowering culm, is a single spikelet of cleistogamous flowers completely surrounded by the leaf sheath. The percentage of cleistogamous flowers produced on any culm depends on the number of chasmogamous and cleistogamous spikelets and the number of flowers in each type of spikelet. All four characters vary among the taxa examined. Danthonia compressa produces, on the average, 50% cleistogamous flowers; D. spicata, 25% cleistogamous flowers; D. sericea and D. epilis, 5% cleistogamous flowers. The degree of cleistogamy in D. spicata is associated with certain habitat features. Populations from mountain sites, disturbed sites, and non-woodland sites produced higher percentages of cleistogamous flowers than did populations from piedmont, undisturbed, and woodland sites, respectively. Grazing may favor increased cleistogamy because cleistogamous flowers are produced lower on the plant. In North Carolina, the most frequently grazed Danthonia taxa also produce the highest percentage of cleistogamous flowers.     

Are pollinators the agents of selection on flower colour and size in irises?

Plant–pollinator interactions are believed to play a major role in the evolution of floral traits. Flower colour and flower size are important for attracting pollinators, directly influencing reproduction, and thus expected to be under pollinator‐mediated selection. Pollinator‐mediated selection is also proposed to play a role in maintaining flower colour polymorphism within populations. However, pigment concentrations, and thus flower colour, are also under selective pressures independent of pollinators. We quantified phenotypic pollinator‐mediated selection on flower colour and size in two colour polymorphic Iris species. Using female fitness, we estimated phenotypic selection on flower colour and size, and tested for pollinator‐mediated selection by comparing selection gradients between flowers open to natural pollination and supplementary pollinated flowers. In both species, we found evidence for pollen limitation, which set the base for pollinator‐mediated selection. In the colour dimorphic Iris lutescens, while pigment concentration and flower size were found to be under selection, this was independent of pollinators. For the polymorphic Iris pumila, pigment concentration is under selective pressure by pollinators, but only for one colour morph. Our results suggest that pollinators are not the main agents of selection on floral traits in these irises, as opposed to the accepted paradigm on floral evolution. This study provides an opposing example to the largely‐accepted theory that pollinators are the major agent of selection on floral traits.     

A generalised mimicry system involving angiosperm flower colour, pollen and bumblebees’ innate colour preferences

Flower colour is a major advertisement signal of zoophilous plants for pollinators. Bees, the main pollinators, exhibit innate colour preferences, which have often been attributed to only one single floral colour, though most flowers display a pattern of two or several colours. The existing studies of floral colour patterns are mostly qualitative studies. Using a model of bee colour vision we quantitatively investigate two questions: whether or not component colours of floral colour patterns may mimic pollen signals, and whether or not bumblebees exhibit innate preferences for distinct parameters of naturally existing floral colour patterns. We analysed the spectral reflectances of 162 plant species with multicoloured flowers and inflorescences, distiniguishing between inner and outer colours of floral colour patterns irrespective of the particular structures so coloured.We found that:– The inner colour of radially symmetrical flowers and inflorescences and of zygomorphic flowers appears less diverse to bees than the peripheral colour.– The inner colour of most radial flowers and inflorescences as well as the inner colour of a large number of non-related zygomorphic flowers appears to bees to be very similar to that of pollen.– Bumblebees (Bombus terrestris) exhibit innate preferences for two-coloured over single-coloured dummy flowers in a spontaneous choice test.– Bumblebees exhibit innate preferences for dummy flowers with a large over those with a small centre area.– Bumblebees exhibit innate preferences for dummy flowers with a centre colour similar to that of pollen over those with another centre colour.Our findings support the hypotheses that the inner component of floral colour patterns could be interpreted as a generalised and little recognised form of mimicry of the colour of visually displayed pollen, that bumblebees exhibit innate preferences regarding colour and size parameters of floral colour patterns, and that these correspond to visually displayed pollen. These findings together suggest a prominent role of floral colour patterns in advertisement to and guidance of naive flower visitors.     

Pollination by hummingbirds and bees in eight syntopic species and a putative hybrid of Ericaceae in Southeastern Brazil

We report on flowering phenology, floral morphology, pollinators, and nectar for eight species and a putative natural hybrid belonging to Agarista, Gaultheria and Gaylussacia that occur syntopically in a montane area. The campanulate to tubular flowers of eight out of nine Ericaceae taxa are primarily pollinated by either hummingbirds or bees. Flowering overlaps in all species but slight differences of floral shape, colour, and nectar characterize pollination by each pollinator group. Differences in floral traits are not large enough to exclude secondary pollinators. Thus, either the main pollinators of a species belonging to its syndrome, or secondary pollinators of a species belonging to different syndromes, may allow for inter-specific crosses.     

In a world of white,flower colour matters: A white–purple transition signals lack of reward in an alpine Euphrasia

The New Zealand alpine flora displays a range of unusual characteristics compared with other alpine floras, in particular the high frequency of species with small white flowers. The presence of both white and bright purple flowers on the same plant in the New Zealand alpine annual creeping eyebright (Euphrasia dyeri Wettst.) provides an ideal opportunity to investigate the significance of flower colour in an environment where coloured flowers are rare. The relationships among flower age, gender phase, reward availability and petal colour were assessed in natural populations of E. dyeri. The effect of pollination on flower colour was tested using hand pollination of bagged flowers. Direct observations and videos of flowers were used to assess patterns of flower visitation by native and introduced pollinators. Unpollinated white E. dyeri flowers changed from white to purple within 6 days. However, pollination of white flowers triggered a significantly faster colour change, typically within 1–2 days. White flowers had receptive stigmas, large amounts of lipid‐rich pollen and small amounts of nectar, whereas stigmas of purple flowers are not receptive and flowers did not provide pollen or nectar rewards. Flowers were mainly visited by native syrphid flies. Both native syrphids and introduced Bombus bees showed a marked avoidance of purple flowers, tending to preferentially visit white flowers. Our study suggests that flower colour change from white to bright purple in E. dyeri functions to direct pollinators to rewarding, receptive flowers. As many Euphrasia L. species are described as having variably coloured flowers, this mechanism may be more widespread in the genus. Furthermore, our results add to the growing evidence that the dominance of white flowers in the New Zealand alpine is not simply due to a lack of colour discrimination among pollinators.     

Phylogenetic evidence for the evolution of ecological specialization in Timema walking‐sticks

We used phylogenetic and ecological information to study the evolution of host‐plant specialization and colour polymorphism in the genus Timema, which comprises 14 species of walking‐sticks that are subject to strong selection for cryptic coloration on their host‐plants. Phylogenetic analysis indicated that this genus consists of three main lineages. Two of the lineages include highly generalized basal species and relatively specialized distal species, and one of the lineages comprises four specialized species. We tested for phylogenetic conservatism in the traits studied via randomizing host‐plant use, and the four basic Timema colour patterns, across the tips of the phylogeny, and determining if the observed number of inferred changes was significantly low compared to the distribution of numbers of inferred changes expected under the null model. This analysis showed that (1) host‐plant use has evolved nonrandomly, such that more closely related species tend to use similar sets of hosts and (2) colour pattern evolution exhibits considerable lability. Inference of ancestral states using maximum parsimony, under four models for the relative ease of gain and loss of plant hosts or colour morphs, showed that (1) for all models with gains of host‐plants even marginally more difficult than losses, and for most optimizations with gains and losses equally difficult, the ancestral Timema were generalized, feeding on the chaparral plants Ceanothus and Adenostoma and possibly other taxa, and (2) for all models with gains of colour morphs more difficult than losses, the ancestral Timema were polymorphic for colour pattern. Generation of null distributions of inferred ancestral states showed that the maximum‐parsimony inference of host‐plant generalization was most robust for the most speciose of the three main Timema lineages. Ancestral states were also inferred using maximum likelihood, after recoding host‐plant use and colour polymorphism as dichotomous characters. Likelihood analyses provided some support for inference of generalization in host‐plant use at ancestral nodes of the two lineages exhibiting mixtures of generalists and specialists, although levels of uncertainty were high. By contrast, likelihood analysis did not estimate ancestral colour morph patterns with any confidence, due to inferred rates of change that were high with respect to speciation rates. Information from biogeography, floristic history and the timing of diversification of the genus are compatible with patterns of inferred ancestral host‐plant use. Diversification in the genus Timema appears to engender three main processes: (1) increased specialization via loss of host‐plants, (2) retention of the same, single, host‐plant and (3) shifts to novel hosts to which lineages were ‘preadapted’ in colour pattern. Our evidence suggests that the radiation of this genus has involved multiple evolutionary transitions from individual‐level specialization (multiple‐niche polymorphism) to population‐level and species‐level specialization. Ecological studies of Timema suggest that such transitions are driven by diversifying selection for crypsis. This paper provides the first phylogeny‐based evidence for the macroevolutionary importance of predation by generalist natural enemies in the evolution of specialization.     

POLYMORPHISM AND PARALLELISM IN THE AFRICAN BUSH-SHRIKES OF THE GENUS MALACONOTUS (INCLUDING CHLOROPHONEUS)

PART 1 Remarkable parallelism exists between two groups of shrikes in the genus Malaconotus: (1) small-billed birds (formerly Chlorophoneus) comprising the polymorphic species multicolor and nigrifrons, the dimorphic olivaceus and the monomorphic sulfureopectus; (2) large-billed birds (formerly Malaconotus sensu stricto)—of the species blanchoti, cruentus, lagdeni, gladiator and alius (and including monteiri). Characters and ranges of the various species are set out and parallelisms between the two groups are listed. M. monteiri is thought to be a colour phase of cruentus rather than a race of blanchoti. It is suggested that the parallelisms are due to the two groups having derived from a common ancestor and that the existing “Malaconotus”, which are monomorphic, may derive directly from polymorphic ancestral stock. part 2 The polymorphism in “Chlorophoneus” can be accounted for by three genes with two alleles each, all interacting with polygenic systems. The homology of the morphs with the colour patterns of monomorphic populations is discussed, and their distributions in the two groups summarised. The interaction of polymorphism with continuous variation in determining the proportion of red to yellow carotenoid is discussed. The evidence bearing upon the status of monteiri is considered. The difference in the incidence of polymorphism between the groups is considered in the light of the analysis into dimorphisms. Some suggestions are made on the significance of parallels between the groups.     

The spatial distribution of nonrewarding artificial flowers affects pollinator attraction

Many species of orchids that do not offer food rewards to pollinators bloom in clusters, early in the season, and are polymorphic for corolla colour. Previous studies suggest that the foraging behaviour of insect pollinators may select for early blooming and colour polymorphism. I tested whether pollinator behaviour can also favour aggregated flowering in these species, in a two-stage laboratory experiment on na?ve bumblebees, Bombus terrestris (L.). In the first stage, the bees were allowed to forage on three colours of artificial flowers that contained sucrose rewards. In the second stage, I added nonrewarding flowers of a fourth colour and recorded the bees' visits to them. The four types of artificial flowers were either arranged in spatially distinct clusters, or were randomly intermingled. I used two reward schedules for each spatial arrangement: constant refilling of reward-containing flowers and probabilistic refilling. Bees that foraged on clustered flowers flew more often to the nonrewarding patch, and made more visits to nonrewarding flowers, than bees that foraged on intermingled flowers. This tendency was obtained both in the constant reward and in the probabilistic reward schedules. The results support the hypothesis that pollinator attraction may select for clustered, synchronized blooming in flowers that do not contain nectar and pollen rewards. Copyright 2000 The Association for the Study of Animal Behaviour.     

Bumblebee-pollination and temporal change of the calyx tube length in Clematis stans(Ranunculaceae)

Clematis stans is dioecious semi-arboreal, with pale purple–blue, nodding, tubulous flowers in a paniculate inflorescence. Both male and female flowers produce nectar from the base of the calyx tube during a flowering period of 3 or 4 days, and are pollinated by two bumblebee species, Bombus diversus and B. honshuensis, with different proboscis lengths. When the flowers open, four sepals constructing a calyx tube separate at the top and their respective tips gradually curl up, so that a tubular part shortens. Observations at two field sites showed that B. diversus (with a longer proboscis) most often visits the flowers with a longer calyx tube, and B. honshuensis (with a shorter proboscis) the flowers with a shorter calyx tube, i.e., later in the flowering period. By changing the calyx tube length, the flowers of C. stans accept the two bumblebee species with different proboscis length as pollinators and thus increase the chance of pollination for each flower. It was also found that the two bumblebee species prefer the male flowers to the female flowers, although the female flowers secrete more nectar as a reward than male flowers. This is likely because they visit the male flowers to collect pollen grains in addition to nectar. Electronic Publication     

Spatial patterns of flower colour variation in native and introduced ranges of Convolvulus arvensis (Convolvulaceae) revealed by citizen-science data and machine learning

• Flower colour polymorphism refers to the presence of multiple colour variants in plant populations. Investigation of this phenomenon led to multiple discoveries, including the principles of heredity and the foundations of population genetics. I examined flower colour variation across native and introduced ranges of Convolvulus arvensis, which exhibits flower colour polymorphism (individuals have white or pink petals).
• To study flower colour variation of this species throughout large geographic scale, I used observations gathered from the iNaturalist platform. To handle a large amount of data, I trained a neural network to classify the plants' morphs based on photographs. After which I performed spatial analyses to examine the patterns of the colour frequency, also in relation to environmental factors.
• The results show that flower colours are polymorphic across the whole species range, but the frequency of pink versus white flowers varies. In the Palearctic, I observed geographic clines of colour morph frequencies: a higher frequency of the pink morph in populations from Northwest Europe, whereas in South and East Europe, towards the eastern edge of the range, the white morph was dominant. In contrast, pattern of colour distribution in North America (where the species is invasive) seems random, but the model indicates a link between higher proportions of pink morphs in mild and humid climates.
• The mechanisms behind the observed patterns remain largely unknown, as changes in a morphs' frequency are not strongly linked to abiotic factors. To understand the spatial pattern, a detailed investigation, accounting for the species' phylogeography is needed. This study provides another example of how the general public may collect data relevant to ecological studies, even when the data are not collected for a specific project.

     

THE IMPACT OF A FLOWER-COLOR POLYMORPHISM ON MATING PATTERNS IN EXPERIMENTAL POPULATIONS OF WILD RADISH (RAPHANUS RAPHANISTRUM L.)

We conducted field experiments to determine how a naturally occurring petal-color polymorphism influences mating patterns in wild radish (Raphanus raphanistrum). The polymorphism is controlled at a single genetic locus, with white petal color being completely dominant to yellow. In experimental populations with equal numbers of yellow- and white-flowered homozygous individuals, insect visitors strongly discriminated against white flowers. Pieris rapae, the most frequent pollinator, was almost 50% more likely to visit yellow than white flowers. Maternal fecundity did not differ between the morphs and was not significantly influenced by a plant's compatibility with potential donors, suggesting that seed production was not limited by receipt of compatible pollen. In contrast, the yellow-flowered morph sired approximately 75% of all seeds produced during the study. This paternity proportion was consistently greater than that expected on the basis of postpollination compatibility measures and was indistinguishable from that expected on the basis of pollinator-visitation frequency. We conclude that the male-fitness advantage of the yellow morph resulted from enhanced pollen export due to the greater attractiveness of its flowers to insect pollinators. With color morphs evenly distributed in experimental arrays, insects did not move assortatively on the basis of petal color, and we found no evidence for assortative pollen flow due to the floral polymorphism. Once postpollination compatibility relationships within populations were taken into account, paternal success of yellow donors did not differ between yellow- and white-flowered maternal plants.     

Selection for individual recognition and the evolution of polymorphic identity signals in Polistes paper wasps

Individual recognition (IR) requires individuals to uniquely identify their social partners based on phenotypic variation. Because IR is so specific, distinctive phenotypes that stand out from the crowd facilitate efficient recognition. Over time, the benefits of unique appearances are predicted to produce a correlation between IR and phenotypic variation. Here, we test whether there is an association between elevated phenotypic polymorphism and IR in paper wasps. Previous work has shown that Polistes fuscatus use variable colour patterns for IR. We test whether two less variable wasp species, Polistes dominulus and Polistes metricus, are capable of IR. As predicted, neither species is capable of IR, suggesting that highly variable colour patterns are confined to Polistes species with IR. This association suggests that elevated phenotypic variation in taxa with IR may be the result of selection for identity signals rather than neutral processes. Given that IR is widespread among social taxa, selection for identity signalling may be an underappreciated mechanism for the origin and maintenance of polymorphism.     

Evidence for rodent pollination in Erica hanekomii (Ericaceae)

Pollination by rodents, originally discovered in Cape Proteaceae, is known from a handful of plant lineages and may be more widespread than current data suggest. Here, we demonstrate the occurrence of this mode of pollination in Erica, the largest plant genus in the Cape flora of South Africa. The localized endemic Erica hanekomii appears to be adapted for pollination by rodents, on account of its low, mat‐forming habit with many dull‐coloured, pendulous flower heads, large volumes (up to c. 30 µL per inflorescence) of sucrose‐dominated nectar, hook‐shaped styles and late winter flowering phenology. Rodents trapped in the vicinity of E. hanekomii plants had many thousands of Erica tetrads in their droppings. Nocturnal laboratory observation showed that Acomys subspinosus (Cape spiny mouse) is adept at manipulating E. hanekomii inflorescences and lapping nectar without destroying gynoecia. During terrarium experiments, pollen was deposited mainly on the rostrum, but also on the chest area and forelegs of A. subspinosus. No birds or insects were seen to visit E. hanekomii in the field. Whereas natural fruit set was 73%, inflorescences from which rodents, but not insects, were excluded using mesh cages, had fruit set of just 3.3%. Exclusion of all visitors resulted in zero seed production. Broken anther rings (a sign of flower visitation in Erica) were observed in 90.1% of flowers in the field and 92% of flowers exposed to rodents in the laboratory, but in virtually none of the flowers from which rodents were selectively excluded. This first record of a non‐flying mammal pollination syndrome in Ericaceae raises interesting new questions about the origins of the rodent pollination system and the selective factors that lead to its evolution. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 163–170.     

Plant phenology and seed predation: interactions between gorses and weevils in Brittany (France)

Exapion ulicis (Forster) and Exapion lemovicinum (Hoffmann) (Coleoptera: Curculionoidea: Apionidae) are seed predators of the three gorse species occurring in Brittany (France): Ulex europaeus L., Ulex gallii Planch., and Ulex minor Roth.(Fabaceae). Host‐plant phenology plays a major role in the relationship between apionid weevils and their gorse species, because larvae develop within gorse pods and adults have to wait for pod dehiscence to be released. We monitored flowering and fruiting phenology of gorse species, weevil reproductive behaviour, and egg‐laying patterns in six natural populations in the native area of these gorse species. At each site, U. europaeus, which flowers mainly in spring, was sympatric with one of two autumn flowering gorse species, U. gallii and U. minor. We noticed that E. ulicis laid eggs in spring and was restricted to U. europaeus whereas E. lemovicinum laid eggs in autumn and was restricted to the two autumn‐flowering species U. gallii and U. minor. Therefore, host specificity depended on gorse phenology, and not on geographic proximity. In addition, the infested pod content showed that E. ulicis laid several eggs per pod and suggested that females chose pods with the highest numbers of seeds. In contrast, E. lemovicinum laid a single egg per pod and showed no preference for pods with many seeds. Finally, the impact of seed predation by E. ulicis was higher than that of E. lemovicinum.     

Remarkable life history polymorphism may be evolving under divergent selection in the silverleaf sunflower

Substantial intraspecific variation in life history is rare and potentially a signal of incipient ecological speciation, if variation is driven by geographically heterogenous natural selection. We present the first report of extensive life history polymorphism in Helianthus argophyllus, the silverleaf sunflower, and examine evidence for its evolution by divergent selection. In 18 populations sampled from across the species range and grown in a common garden, most quantitative traits covaried such that individuals could be assigned to two distinct life history syndromes: tall and late flowering with small initial flowerheads, or short and early flowering with larger initial flowerheads. Helianthus argophyllus exhibits regional genetic structure, but this population structure does not closely correspond with patterns of phenotypic variation. The early‐flowering syndrome is primarily observed in populations from coastal barrier islands, while populations from the nearby mainland coast, although geographically and genetically close, are primarily late flowering. Additionally, several traits are more differentiated among regions than expected based on neutral genetic divergence (Q_ST > F_ST), including the first principal component score corresponding with life history syndrome. This discordance between patterns of phenotypic and genetic variation suggests that divergent selection is driving genetic differences in life history across the species range. If so, the silverleaf sunflower may be in early stages of ecological speciation.     

Small but attractive: female‐biased nectar production and floral visitors in a dimorphic shrub

• In sexually dimorphic species, hermaphrodite flowers in gynodioecious species or male flowers in dioecious species are often larger and produce more nectar than their conspecific female flowers. As a consequence, hermaphrodite or male flowers frequently receive more pollinator visits.
• Sex ratio, flower size, floral display, nectar production and floral visits were evaluated in two natural populations of Fuchsia thymifolia, a morphologically gynodioecious but functionally subdioecious insect‐pollinated shrub.
• Sex ratio did not differ from the expected 1:1 in the two studied populations. As expected, hermaphrodite flowers were larger than female flowers, but in contrast to the general pattern, hermaphrodite flowers did not produce nectar or produced much less than female flowers. Flower visitors were flies (68%) and bumblebees (24%), both of which showed a preference for female flowers. No sex difference was detected in either flower longevity or floral display across the flowering season.
• Higher nectar production by females may attract more pollinators, and may be a strategy to enhance female reproductive success in this species. Finally, floral dimorphism and insect preferences did not seem to hamper the maintenance of sub‐dioecy or prevent the evolution of dioecy in F. thymifolia.

     

Variation in the phenology and abundance of flowering by native and exotic plants in subalpine meadows

The timing and abundance of flower production is important to the reproductive success of angiosperms as well as pollinators and floral and seed herbivores. Exotic plants often compete with native plants for space and limiting resources, potentially altering community floral dynamics. We used observations and a biomass-removal experiment to explore the effects of an invasive exotic flowering plant, Linaria vulgaris, on community and individual species flowering phenology and abundance in subalpine meadows in Colorado, USA. Invasion by L. vulgaris was associated with a shift in both the timing and abundance of community flowering. Invaded plant communities exhibited depressed flowering by 67% early in the season relative to uninvaded communities, but invaded sites produced 7.6 times more flowers than uninvaded sites once L. vulgaris began flowering. This increase in flowers at the end of the season was driven primarily by prolific flowering of L. vulgaris. We also found lower richness and evenness of resident flowering species in invaded plots during the period of L. vulgaris flowering. At the species level, a common native species (Potentilla pulcherrima) produced 71% fewer flowers in invaded relative to uninvaded plots, and the species had reduced duration of flowering in invaded relative to uninvaded sites. This result suggests that L. vulgaris does not simply alter the flowering of subordinate species but also the flowering of an individual common species in the plant community. We then used observational data to explore the relationship between L. vulgaris density and resident floral production but found only partial evidence that higher densities of L. vulgaris were associated with stronger effects on resident floral production. Taken together, results suggest that a dominant invasive plant can affect community and individual-species flowering.