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1.  Morphofunctional Traits and Pollination Mechanisms of Coronilla emerus L. Flowers (Fabaceae) 
The Scientific World Journal  2012;2012:381575.
It is accepted that the papilionaceous corolla of the Fabaceae evolved under the selective pressure of bee pollinators. Morphology and function of different parts of Coronilla emerus L. flowers were related to their role in the pollination mechanism. The corolla has a vexillum with red nectar lines, a keel hiding stamens and pistil, and two wing petals fasten to the keel with two notched folds. Pollinators land on the complex of keel and wings, trigger the protrusion of pollen and finally of the stigma from the keel tip. Data on pollen viability and stigma receptivity prove that flowers are proterandrous. The results of hand-pollination experiments confirmed that insects are fundamental to set seed. Interaction with pollinators allows not only the transport of pollen but also the rupture of the stigmatic cuticle, necessary to achieve both allogamy and autogamy. Field observations showed that Hymenoptera, Lepidoptera, and Diptera visited the flowers. Only some of the Hymenoptera landed on the flowers from the front and elicited pollination mechanisms. Most of the insects sucked the nectar from the back without any pollen transfer. Finally, morphological and functional characteristics of C. emerus flowers are discussed in terms of floral larceny and reduction in pollination efficiency.
PMCID: PMC3361249  PMID: 22666114
2.  Gloss, Colour and Grip: Multifunctional Epidermal Cell Shapes in Bee- and Bird-Pollinated Flowers 
PLoS ONE  2014;9(11):e112013.
Flowers bear the function of filters supporting the attraction of pollinators as well as the deterrence of floral antagonists. The effect of epidermal cell shape on the visual display and tactile properties of flowers has been evaluated only recently. In this study we quantitatively measured epidermal cell shape, gloss and spectral reflectance of flowers pollinated by either bees or birds testing three hypotheses: The first two hypotheses imply that bee-pollinated flowers might benefit from rough surfaces on visually-active parts produced by conical epidermal cells, as they may enhance the colour signal of flowers as well as the grip on flowers for bees. In contrast, bird-pollinated flowers might benefit from flat surfaces produced by flat epidermal cells, by avoiding frequent visitation from non-pollinating bees due to a reduced colour signal, as birds do not rely on specific colour parameters while foraging. Moreover, flat petal surfaces in bird-pollinated flowers may hamper grip for bees that do not touch anthers and stigmas while consuming nectar and thus, are considered as nectar thieves. Beside this, the third hypothesis implies that those flower parts which are vulnerable to nectar robbing of bee- as well as bird-pollinated flowers benefit from flat epidermal cells, hampering grip for nectar robbing bees. Our comparative data show in fact that conical epidermal cells are restricted to visually-active parts of bee-pollinated flowers, whereas robbing-sensitive parts of bee-pollinated as well as the entire floral surface of bird-pollinated flowers possess on average flat epidermal cells. However, direct correlations between epidermal cell shape and colour parameters have not been found. Our results together with published experimental studies show that epidermal cell shape as a largely neglected flower trait might act as an important feature in pollinator attraction and avoidance of antagonists, and thus may contribute to the partitioning of flower-visitors.
PMCID: PMC4219824  PMID: 25369510
3.  Sexual Dimorphism of Staminate- and Pistillate-Phase Flowers of Saponaria officinalis (Bouncing Bet) Affects Pollinator Behavior and Seed Set 
PLoS ONE  2014;9(4):e93615.
The sequential separation of male and female function in flowers of dichogamous species allows for the evolution of differing morphologies that maximize fitness through seed siring and seed set. We examined staminate- and pistillate-phase flowers of protandrous Saponaria officinalis for dimorphism in floral traits and their effects on pollinator attraction and seed set. Pistillate-phase flowers have larger petals, greater mass, and are pinker in color, but due to a shape change, pistillate-phase flowers have smaller corolla diameters than staminate-phase flowers. There was no difference in nectar volume or sugar content one day after anthesis, and minimal evidence for UV nectar guide patterns in staminate- and pistillate-phase flowers. When presented with choice arrays, pollinators discriminated against pistillate-phase flowers based on their pink color. Finally, in an experimental garden, in 2012 there was a negative correlation between seed set of an open-pollinated, emasculated flower and pinkness (as measured by reflectance spectrometry) of a pistillate-phase flower on the same plant in plots covered with shade cloth. In 2013, clones of genotypes chosen from the 2012 plants that produced pinker flowers had lower seed set than those from genotypes with paler flowers. Lower seed set of pink genotypes was found in open-pollinated and hand-pollinated flowers, indicating the lower seed set might be due to other differences between pink and pale genotypes in addition to pollinator discrimination against pink flowers. In conclusion, staminate- and pistillate-phase flowers of S. officinalis are dimorphic in shape and color. Pollinators discriminate among flowers based on these differences, and individuals whose pistillate-phase flowers are most different in color from their staminate-phase flowers make fewer seeds. We suggest morphological studies of the two sex phases in dichogamous, hermaphroditic species can contribute to understanding the evolution of sexual dimorphism in plants without the confounding effects of genetic differences between separate male and female individuals.
PMCID: PMC3972141  PMID: 24690875
4.  Keel petal incision: a simple and efficient method for genetic crossing in Medicago truncatula 
Plant Methods  2014;10:11.
Genetic crossing is an essential tool in both forward and reverse genetic approaches to understand the biological functions of genes. For Medicago truncatula (barrel medic) various crossing techniques have been used which differ in the methods used to dissect the female parent’s unopened flower bud to remove immature anthers for prevention of self-pollination. Previously described methods including front, side or back incision methods may damage the flower bud, impeding successful fertilization and/or seed development because they may allow pollen to dislodge and floral organs to desiccate after crossing, all of which diminish the success rates of crossing.
We report the keel petal incision method for genetic crossing in M. truncatula ecotype R108 and demonstrate successful crosses with two other M. truncatula ecotypes, A17 and A20. In the method presented here, an incision is made along the central line of the keel petal from the bottom 1/3rd of the female parent’s flower bud to its distal end. This allows easy removal of anthers from the flower bud and access for cross-pollination. After pollination, the stigma and the deposited pollen from the male donor are covered by the keel petal, wing petals and standard petal, forming a natural pouch. The pouch prevents dislodging of deposited pollen from the stigma and protects the internal floral organs from drying out, without using cling-film or water-containing chambers to maintain a humid environment. The keel petal incision method showed an approximate 80% success rate in the M. truncatula R108 ecotype and also in other ecotypes including Jemalong A17 and A20.
Our keel petal incision protocol shows marked improvement over existing methods with respect to the ease of crossing and the percentage of successful crosses. Developed for the M. truncatula R108 ecotype, the protocol has been demonstrated with A17 and A20 ecotypes and is expected to work with other ecotypes. Investigators of varying experience have achieved genetic crosses in M. truncatula using this method.
PMCID: PMC4070640  PMID: 24966878
Legume; Genetic crossing; Barrel medic; Medicago truncatula; Artificial hybridization; Keel petal
5.  A Regulatory Network for Coordinated Flower Maturation 
PLoS Genetics  2012;8(2):e1002506.
For self-pollinating plants to reproduce, male and female organ development must be coordinated as flowers mature. The Arabidopsis transcription factors AUXIN RESPONSE FACTOR 6 (ARF6) and ARF8 regulate this complex process by promoting petal expansion, stamen filament elongation, anther dehiscence, and gynoecium maturation, thereby ensuring that pollen released from the anthers is deposited on the stigma of a receptive gynoecium. ARF6 and ARF8 induce jasmonate production, which in turn triggers expression of MYB21 and MYB24, encoding R2R3 MYB transcription factors that promote petal and stamen growth. To understand the dynamics of this flower maturation regulatory network, we have characterized morphological, chemical, and global gene expression phenotypes of arf, myb, and jasmonate pathway mutant flowers. We found that MYB21 and MYB24 promoted not only petal and stamen development but also gynoecium growth. As well as regulating reproductive competence, both the ARF and MYB factors promoted nectary development or function and volatile sesquiterpene production, which may attract insect pollinators and/or repel pathogens. Mutants lacking jasmonate synthesis or response had decreased MYB21 expression and stamen and petal growth at the stage when flowers normally open, but had increased MYB21 expression in petals of older flowers, resulting in renewed and persistent petal expansion at later stages. Both auxin response and jasmonate synthesis promoted positive feedbacks that may ensure rapid petal and stamen growth as flowers open. MYB21 also fed back negatively on expression of jasmonate biosynthesis pathway genes to decrease flower jasmonate level, which correlated with termination of growth after flowers have opened. These dynamic feedbacks may promote timely, coordinated, and transient growth of flower organs.
Author Summary
Perfect flowers have both male organs that produce and release pollen and female organs that make and harbor seeds. Flowers also often attract pollinators using visual or chemical signals. So that male, female, and pollinator attraction functions occur at the right time, flower organs must grow and mature in a coordinated fashion. In the model self-pollinating plant Arabidopsis, a transcriptional network regulates genes that ensure coordinated growth of different flower organs, as well as pollen release and gynoecium (female) competence to support pollination. This network also regulates nectary development and production of volatile chemicals that may attract or repel insects. We have studied growth, chemical signal levels, and gene expression in mutants affected in components of this network, in order to determine how flower growth is controlled. Several plant hormones act in a cascade that promotes flower maturation. Moreover, regulatory feedback loops affect the timing and extent of developmental steps. Positive feedbacks may ensure that the development of different flower organs is coordinated and rapid, whereas negative feedbacks may allow growth to cease once flowers have opened. Our results provide a framework to understand how flower opening and reproduction are coordinated in Arabidopsis and other flowering plants.
PMCID: PMC3276552  PMID: 22346763
6.  Variation in highbush blueberry floral volatile profiles as a function of pollination status, cultivar, time of day and flower part: implications for flower visitation by bees 
Annals of Botany  2011;107(8):1377-1390.
Background and Aims
Studies of the effects of pollination on floral scent and bee visitation remain rare, particularly in agricultural crops. To fill this gap, the hypothesis that bee visitation to flowers decreases after pollination through reduced floral volatile emissions in highbush blueberries, Vaccinium corymbosum, was tested. Other sources of variation in floral emissions and the role of floral volatiles in bee attraction were also examined.
Pollinator visitation to blueberry flowers was manipulated by bagging all flowers within a bush (pollinator excluded) or leaving them unbagged (open pollinated), and then the effect on floral volatile emissions and future bee visitation were measured. Floral volatiles were also measured from different blueberry cultivars, times of the day and flower parts, and a study was conducted to test the attraction of bees to floral volatiles.
Key Results
Open-pollinated blueberry flowers had 32 % lower volatile emissions than pollinator-excluded flowers. In particular, cinnamyl alcohol, a major component of the floral blend that is emitted exclusively from petals, was emitted in lower quantities from open-pollinated flowers. Although, no differences in cinnamyl alcohol emissions were detected among three blueberry cultivars or at different times of day, some components of the blueberry floral blend were emitted in higher amounts from certain cultivars and at mid-day. Field observations showed that more bees visited bushes with pollinator-excluded flowers. Also, more honey bees were caught in traps baited with a synthetic blueberry floral blend than in unbaited traps.
Greater volatile emissions may help guide bees to unpollinated flowers, and thus increase plant fitness and bee energetic return when foraging in blueberries. Furthermore, the variation in volatile emissions from blueberry flowers depending on pollination status, plant cultivar and time of day suggests an adaptive role of floral signals in increasing pollination of flowers.
PMCID: PMC3101143  PMID: 21498566
Vaccinium corymbosum; honey bees; bumble bees; volatile organic compounds; diurnal rhythm; nectar production; site of emission
7.  Spatial variation in selection on corolla shape in a generalist plant is promoted by the preference patterns of its local pollinators 
An adaptive role of corolla shape has been often asserted without an empirical demonstration of how natural selection acts on this trait. In generalist plants, in which flowers are visited by diverse pollinator fauna that commonly vary spatially, detecting pollinator-mediated selection on corolla shape is even more difficult. In this study, we explore the mechanisms promoting selection on corolla shape in the generalist crucifer Erysimum mediohispanicum Polatschek (Brassicaceae). We found that the main pollinators of E. mediohispanicum (large bees, small bees and bee flies) discriminate between different corolla shapes when offered artificial flowers without reward. Importantly, different pollinators prefer different shapes: bees prefer flowers with narrow petals, whereas bee flies prefer flowers with rounded overlapping petals. We also found that flowers with narrow petals (those preferred by bees) produce both more pollen and nectar than those with rounded petals. Finally, different plant populations were visited by different faunas. As a result, we found spatial variation in the selection acting on corolla shape. Selection favoured flowers with narrow petals in the populations where large or small bees are the most abundant pollinator groups. Our study suggests that pollinators, by preferring flowers with high reward, exert strong selection on the E. mediohispanicum corolla shape. The geographical variation in the pollinator-mediated selection on E. mediohispanicum corolla shape suggests that phenotypic evolution and diversification can occur in this complex floral trait even without specialization.
PMCID: PMC2603243  PMID: 18544510
corolla shape evolution; pollinator preference; spatial variation; geometric morphometrics; nectar; pollen
8.  Pollinators of the Rocky Mountain columbine: temporal variation, functional groups and associations with floral traits 
Annals of Botany  2009;103(9):1567-1578.
Background and Aims
Pollinators together with other biotic and some abiotic factors can select for floral traits. However, variation in pollinator abundance over time and space can weaken such selection. In the present study, the variation in pollinator abundance over time and space was examined in populations of the Rocky Mountain columbine. The variation in three floral traits is described and correlations between pollinator type, functional pollinator groups or altitude and floral traits are examined.
Pollinator observations took place in six Aquilegia coerulea populations over 1–4 years and spur length, flower colour and sepal length were measured in 12 populations. Pollinator abundance, measured as visits per flower per hour, was compared among populations and years. Pollinators were grouped into two functional groups: pollen or nectar collectors. The following associations were examined: annual presence of hawkmoths and whiter flowers with longer spurs; the presence of Sphinx vashti and longer spurs; and higher altitudes and whiter flowers. The study looked at whether an increase in the proportion of hawkmoths in a population was associated with whiter and larger flowers with longer spurs.
Key Results
The abundance of different pollinator groups varied over time and space. Floral traits varied among populations. Higher altitude was correlated with bluer flowers. Whiter flowers were associated with the annual presence of hawkmoths. Populations visited by Sphinx vashti had longer spurs than populations visited only by Hyles lineata. Populations with greater percentage of nectar-collecting pollinators did not have whiter, larger flowers with longer spurs.
Despite the large variation in pollinator abundance over time and space, one species of bumble-bee or hawkmoth tended to predominate in each population each year. Future studies of Aquilegia coerulea should examine the specific influences of pollinators and the environment on flower colour and of hawkmoth species on spur length.
PMCID: PMC2701757  PMID: 19414518
Aquilegia coerulea; columbine; pollinator abundance; bumble-bee; hawkmoth, flower colour; spur length; functional pollinator group; altitude; floral trait
9.  Extreme variation in floral characters and its consequences for pollinator attraction among populations of an Andean cactus 
Annals of Botany  2009;103(9):1489-1500.
Background and aims
A South American cactus species, Echinopsis ancistrophora (Cactaceae), with dramatic among-population variation in floral traits is presented.
Eleven populations of E. ancistrophora were studied in their habitats in northern Argentina, and comparisons were made of relevant floral traits such as depth, stigma position, nectar volume and sugar concentration, and anthesis time. Diurnal and nocturnal pollinator assemblages were evaluated for populations with different floral trait combinations.
Key Results
Remarkable geographical variations in floral traits were recorded among the 11 populations throughout the distribution range of E. ancistrophora, with flower lengths ranging from 4·5 to 24·1 cm. Other floral traits associated with pollinator attraction also varied in a population-specific manner, in concert with floral depth. Populations with the shortest flowers showed morning anthesis and those with the longest flowers opened at dusk, whereas those with flowers of intermediate length opened at unusual times (2300–0600 h). Nectar production varied non-linearly with floral length; it was absent to low (population means up to 15 µL) in short- to intermediate-length flowers, but was high (population means up to 170 µL) in the longest tubed flowers. Evidence from light-trapping of moths, pollen carriage on their bodies and moth scale deposition on stigmas suggests that sphingid pollination is prevalent only in the four populations with the longest flowers, in which floral morphological traits and nectar volumes match the classic expectations for the hawkmoth pollination syndrome. All other populations, with flowers 4·5–15 cm long, were pollinated exclusively by solitary bees.
The results suggest incipient differentiation at the population level and local adaptation to either bee or hawkmoth (potentially plus bee) pollination.
PMCID: PMC2701769  PMID: 19342397
Pollination; floral biology; Echinopsis ancistrophora; cactus; Cactaceae; hawkmoth; bee
10.  Are there pollination syndromes in the Australian epacrids (Ericaceae: Styphelioideae)? A novel statistical method to identify key floral traits per syndrome 
Annals of Botany  2013;112(1):141-149.
Background and Aims
Convergent floral traits hypothesized as attracting particular pollinators are known as pollination syndromes. Floral diversity suggests that the Australian epacrid flora may be adapted to pollinator type. Currently there are empirical data on the pollination systems for 87 species (approx. 15 % of Australian epacrids). This provides an opportunity to test for pollination syndromes and their important morphological traits in an iconic element of the Australian flora.
Data on epacrid–pollinator relationships were obtained from published literature and field observation. A multivariate approach was used to test whether epacrid floral attributes related to pollinator profiles. Statistical classification was then used to rank floral attributes according to their predictive value. Data sets excluding mixed pollination systems were used to test the predictive power of statistical classification to identify pollination models.
Key Results
Floral attributes are correlated with bird, fly and bee pollination. Using floral attributes identified as correlating with pollinator type, bird pollination is classified with 86 % accuracy, red flowers being the most important predictor. Fly and bee pollination are classified with 78 and 69 % accuracy, but have a lack of individually important floral predictors. Excluding mixed pollination systems improved the accuracy of the prediction of both bee and fly pollination systems.
Although most epacrids have generalized pollination systems, a correlation between bird pollination and red, long-tubed epacrids is found. Statistical classification highlights the relative importance of each floral attribute in relation to pollinator type and proves useful in classifying epacrids to bird, fly and bee pollination systems.
PMCID: PMC3690994  PMID: 23681546
Epacridaceae (epacrids); Ericaceae; multivariate analysis; plant–pollinator interactions; pollination syndromes; Random Forests; statistical classification; Styphelioideae
11.  Functional morphology and wasp pollination of two South American asclepiads (Asclepiadoideae–Apocynaceae) 
Annals of Botany  2011;109(1):77-93.
Background and Aims
The extreme complexity of asclepiad flowers (Asclepiadoideae–Apocynaceae) has generated particular interest in the pollination biology of this group of plants especially in the mechanisms involved in the pollination processes. This study compares two South American species, Morrenia odorata and Morrenia brachystephana, with respect to morphology and anatomy of flower structures, dynamic aspects of the pollination mechanism, diversity of visitors and effectiveness of pollinators.
Floral structure was studied with fresh and fixed flowers following classical techniques. The pollination mechanism was studied by visiting fresh flowers in the laboratory with artificial pollinator body parts created with an eyelash. Morphometric and nectar measurements were also taken. Pollen transfer efficiency in the flowers was calculated by recording the frequency of removed and inserted pollinia. Visitor activity was recorded in the field, and floral visitors were captured for subsequent analysis of pollen loads. Finally, pollinator effectiveness was calculated with an index.
Key Results
The detailed structure of the flowers revealed a complex system of guide rails and chambers precisely arranged in order to achieve effective pollinaria transport. Morrenia odorata is functionally specialized for wasp pollination, and M. brachystephana for wasp and bee pollination. Pollinators transport chains of pollinaria adhered to their mouthparts.
Morrenia odorata and M. brachystephana present differences in the morphology and size of their corona, gynostegium and pollinaria, which explain the differences in details of the functioning of the general pollination mechanism. Pollination is performed by different groups of highly effective pollinators. Morrenia species are specialized for pollination mainly by several species of wasps, a specialized pollination which has been poorly studied. In particular, pompilid wasps are reported as important pollinators in other regions outside South Africa. A putative new function of nectar in asclepiads is presented, as it would be contributing to the pollination mechanism.
PMCID: PMC3241588  PMID: 22025522
Morrenia odorata; Morrenia brachystephana; asclepiads; functional morphology; pollination mechanism; wasps; pollinator effectiveness
12.  Floral dimorphism in plant populations with combined versus separate sexes 
Annals of Botany  2011;108(4):765-776.
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.
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.
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.
PMCID: PMC3170149  PMID: 21385777
Sexual dimorphism; flower size; daily floral display; sexual selection; sex ratios; monoecy; dioecy; Sagittaria latifolia
13.  Evolutionary consequences of shifts to bird-pollination in the Australian pea-flowered legumes (Mirbelieae and Bossiaeeae) 
Interactions with pollinators are proposed to be one of the major drivers of diversity in angiosperms. Specialised interactions with pollinators can lead to specialised floral traits, which collectively are known as a pollination syndrome. While it is thought that specialisation to a pollinator can lead to either an increase in diversity or in some cases a dead end, it is not well understood how transitions among specialised pollinators contribute to changes in diversity. Here, we use evolutionary trait reconstruction of bee-pollination and bird-pollination syndromes in Australian egg-and-bacon peas (Mirbelieae and Bossiaeeae) to test whether transitions between pollination syndromes is correlated with changes in species diversity. We also test for directionality in transitions that might be caused by selection by pollinators or by an evolutionary ratchet in which reversals to the original pollination syndrome are not possible.
Trait reconstructions of Australian egg-and-bacon peas suggest that bee-pollination syndrome is the ancestral form and that there has been replicated evolution of bird-pollination syndromes. Reconstructions indicate potential reversals from bird- to bee-pollination syndromes but this is not consistent with morphology. Species diversity of bird-pollination syndrome clades is lower than that of their bee-pollination syndrome sisters.
We estimated the earliest transitions from bee- to bird-pollination syndrome occurred between 30.8 Ma and 10.4 Ma. Geographical structuring of pollination syndromes was found; there were fewer bird-pollination species in the Australian southeast temperate region compared to other regions of Australia.
A consistent decrease in diversification rate coincident with switches to bird pollination might be explained if greater dispersal by bird pollinators results in higher levels of connectivity among populations and reduced chances of allopatric speciation.
The earliest transitions overlap with the early diversification of Australian honeyeaters – the major lineage of pollinating birds in Australia. Our findings are consistent with the idea that environment and availability of pollinators are important in the evolution of pollination syndromes. Changes in flower traits as a result of transitions to bird-pollination syndrome might also limit reversals to a bee-pollination syndrome.
PMCID: PMC4015313  PMID: 24602227
Pollination syndrome; Adaptive radiation; Ancestral state reconstruction; Diversification
14.  Wild bees preferentially visit Rudbeckia flower heads with exaggerated ultraviolet absorbing floral guides 
Biology Open  2014;3(3):221-230.
Here, we report on the results of an experimental study that assessed the visitation frequency of wild bees to conspecific flowers with different sized floral guides. UV absorbent floral guides are ubiquitous in Angiosperms, yet surprisingly little is known about conspecific variation in these guides and very few studies have evaluated pollinator response to UV guide manipulation. This is true despite our rich understanding about learning and color preferences in bees. Historical dogma indicates that flower color serves as an important long-range visual signal allowing pollinators to detect the flowers, while floral guides function as close-range signals that direct pollinators to a reward. We initiated the work presented here by first assessing the population level variation in UV absorbent floral guides for conspecific flowers. We assessed two species, Rudbeckia hirta and R. fulgida. We then used several petal cut-and-paste experiments to test whether UV floral guides can also function to attract visitors. We manipulated floral guide size and evaluated visitation frequency. In all experiments, pollinator visitation rates were clearly associated with floral guide size. Diminished floral guides recruited relatively few insect visitors. Exaggerated floral guides recruited more visitors than smaller or average sized guides. Thus, UV floral guides play an important role in pollinator recruitment and in determining the relative attractiveness of conspecific flower heads. Consideration of floral guides is therefore important when evaluating the overall conspicuousness of flower heads relative to background coloration. This work raises the issue of whether floral guides serve as honest indicators of reward, since guide size varies in nature for conspecific flowers at the same developmental stage and since preferences for larger guides were found. To our knowledge, these are the first cut-and-paste experiments conducted to examine whether UV absorbent floral guides affect visitation rates and pollinator preference.
PMCID: PMC4001241  PMID: 24585774
Bee; Rudbeckia; Flower; Pollination; Opsin; Vision; Pollinator
15.  Effect of Pollination on Floral Longevity and Costs of Delaying Fertilization in the Out-crossing Polygala vayredae Costa (Polygalaceae) 
Annals of Botany  2008;102(6):1043-1048.
Background and Aims
The effect of pollination on flower life span has been widely studied, but so far little attention has been paid to the reproductive consequences of delayed pollination in plants with long floral life spans. In the present study, Polygala vayredae was used to answer the following questions. (1) How does male and female success affect the floral longevity of individual flowers? (2) How does delaying fertilization affect the female fitness of this species?
Floral longevity was studied after experimental pollinations involving male and/or female accomplishment, bagging and open pollination. The reproductive costs of a delay in the moment of fertilization were evaluated through fruit set, seed–ovule ratio and seed weight, after pollination of flowers that had been bagged for 2–18 d.
Key Results
Senescence of the flowers of P. vayredae was activated by pollen reception on the stigmatic papillae, while pollen removal had no effect on floral longevity. Nonetheless, a minimum longevity of 8 d was detected, even after successful pollination and pollen dissemination. This period may be involved with the enhancement of male accrual rates, as the female accomplishment is generally achieved after the first visit. Floral life span of open-pollinated flowers was variable and negatively correlated with pollinator visitation rates. Delayed pollination had a major impact on the reproductive success of the plant, with fruit set, seed–ovule ratio and seed weight being significantly diminished with the increase of flower age at the moment of fertilization.
A strong relationship between pollination and floral longevity was observed. Flowers revealed the ability to extend or reduce their longevity, within some limits, in response to the abundance of efficient pollinators (i.e. reproductive fulfilment rates). Furthermore, with scarce or unpredictable pollinators, a long floral life span could maintain the opportunity for fertilization but would also have reproductive costs on production of offspring. Reduced female fitness late in the flower's life could shift the cost–benefit balance towards a shorter life span, partially counteracting the selection for longer floral life span potentially mediated by scarce pollination services.
PMCID: PMC2712401  PMID: 18829587
Delayed pollination; endemic species; flower longevity; life span; pollen limitation; pollination; pollinator scarcity; Polygala vayredae; Polygalaceae; reproductive consequences; secondary pollen presentation
16.  A pollinator shift explains floral divergence in an orchid species complex in South Africa 
Annals of Botany  2013;113(2):277-288.
Background and Aims
Floral diversification driven by shifts between pollinators has been one of the key explanations for the radiation of angiosperms. According to the Grant–Stebbins model of pollinator-driven speciation, these shifts result in morphologically distinct ‘ecotypes’ which may eventually become recognizable as species. The current circumscription of the food-deceptive southern African orchid Eulophia parviflora encompasses a highly variable monophyletic species complex. In this study, two forms were identified within this complex that differ in distribution, floral morphology, scent chemistry and phenology, and a test was made of whether these differences represent adaptations for different pollinators.
Methods and Results
Multivariate analysis of floral and vegetative traits revealed that there are at least two discrete morphological forms in the species complex. Field observations revealed that each form is pollinated by a different insect species, and thus represent distinct ecotypes. The early-flowering coastal form which has long spurs and floral scent dominated by sesquiterpene compounds is pollinated exclusively by the long-tongued bee Amegilla fallax (Apidae, Anthophorinae), while the late-flowering inland form with short spurs and floral scent dominated by benzenoid compounds is pollinated exclusively by the beetle Cyrtothyrea marginalis (Cetoniinae; Scarabaeidae). Choice experiments in a Y-maze olfactometer showed that beetles are preferentially attracted to the scent of the short-spurred form. A spur-shortening experiment showed that long spurs are required for effective pollination of the bee-pollinated form. Although it was initially thought likely that divergence occurred across a geographical pollinator gradient, plants of the long-spurred form were effectively pollinated when transplanted to an inland locality outside the natural coastal range of this form. Thus, the underlying geographical basis for the evolution of ecotypes in the E. parviflora complex remains uncertain, although early flowering in the long-spurred form to exploit the emergence of naïve bees may restrict this form to coastal areas where there is no frost that would damage flower buds. Later flowering of the short-spurred form coincides closely with the emergence of the pollinating beetles following winter frosts.
This study identifies a shift between bee and beetle pollination as the main driver of floral divergence in an orchid species complex. Floral scent and spur length appear to be key traits in mediating this evolutionary transition.
PMCID: PMC3890387  PMID: 24107684
Grant–Stebbins model; pollinator-driven speciation; pollination ecotypes; scent; Eulophia; Orchidaceae; phenology; beetle pollination; Cetoniinae; bee pollination; Y-maze olfactometer; Amegilla
17.  The Pollination Ecology of Paraboea rufescens (Gesneriaceae): a Buzz-pollinated Tropical Herb with Mirror-image Flowers 
Annals of Botany  2006;97(3):371-376.
• Background and Aims Gesneriaceae is a pantropical plant family with over 3000 species. A great variety of pollination mechanisms have been reported for the neotropical members of the family, but the details of buzz-pollination and enantiostyly for the family have not been described. We investigated the floral biology and pollination ecology of Paraboea rufescens in Xishuangbanna, south-west China, considering three aspects: (1) the type of enantiostyly exhibited; (2) whether the species is self-compatible; and (3) whether pollinator behaviour could enhance the precision of pollen transfer between flowers of contrasting stylar orientation.
• Methods Flowering phenology was monitored once a month during vegetative growth, and once a week during flowering both in the field and under cultivation. Pollination manipulations and pollinator observation in the field were conducted.
• Key Results Anthesis occurred early during the morning, and flowers remained open for 1–5 d, depending on weather conditions. Controlled pollinations revealed that P. rufescens is self-compatible, and exhibited inbreeding depression in seed set. Plants were pollinator limited in natural populations. The similar stylar deflection among flowers within a plant limits autonomous self-pollination as well as pollination between flowers. Two species of bumble bees (Bombus spp.), Amegila malaccensis and Nomia sp. effectively pollinated P. rufescens. These pollinators visited flowers in search of pollen with almost the same frequency. None of the pollinators appeared to discriminate between left- or right-handed flowers.
• Conclusions Paraboea rufescens exhibits monomorphic enantiostylous flowers and a buzz-pollination syndrome. Floral morphology in P. rufescens and pollinator foraging behaviour seems likely to reduce self-pollination and pollinations between flowers of the same stylar deflection.
PMCID: PMC2803634  PMID: 16371444
Buzz-pollination; enantiostyly; Gesneriaceae; mirror-image flowers; Paraboea rufescens; reproductive biology; Xishuangbanna
18.  The reproductive strategy of a pollinator-limited Himalayan plant, Incarvillea mairei (Bignoniaceae) 
BMC Plant Biology  2013;13:195.
Plants may adapt to alpine habitats by specialization in the reproductive strategy and functional aspects of their flowers and pollination systems. Alpine habitats reduce the opportunities for cross-pollination in a relatively high proportion of alpine plant species, and self-pollination may be favored in these adverse conditions. Here, we investigated the mating system and pollination of Incarvillea mairei, a perennial Himalayan herb typically found at altitudes between 3000 and 4500 m.
Analyses of floral morphology, observation of plant-pollinator interactions, and hand pollination experiments were conducted in three natural populations. Outcrossing rates and effective numbers of pollen donors were assessed in 45 open-pollinated families by using progeny analysis based on seven microsatellite markers. Incarvillea mairei displayed a set of apparently specialized floral traits, the stigma is sensitive to touch and close immediately and its reopening allows a second opportunity for the receipt of pollen. The species is fully self-compatible but employs a predominantly outcrossing mating system according to parentage analysis (tm > 0.9). Fruit set was low (26.3%), whereas seed set was high (67.2%), indicating that this species suffers pollinator limitation. Its main effective pollinator was Halictus sp., and visitation frequency was low.
Floral features such as having a sensitive stigma and anther-prongs, in conjunction with pollinator behavior, function together contributing to a set of unique reproductive adaptations that enhance outcrossing success. The increased floral longevity and high pollination efficiency operated as compensatory mechanisms to counteract low pollinator visitation frequency.
PMCID: PMC4219382  PMID: 24289097
Himalaya-Hengduan mountains region; Incarvillea mairei; Plant-pollinator interaction; Pollinator limitation; Sensitive stigma
19.  Bilabiate Flowers: The Ultimate Response to Bees? 
Annals of Botany  2007;100(2):361-374.
Background and Aims
Bilabiate flowers have evolved in many lineages of the angiosperms, thus representing a convincing example of parallel evolution. Similar to keel blossoms, they have obviously evolved in order to protect pollen against pollen-collecting bees. Although many examples are known, a comprehensive survey on floral diversity and functional constraints of bilabiate flowers is lacking. Here, the concept is widened and described as a general pattern.
The present paper is a conceptional review including personal observations of the authors. To form a survey on the diversity of bilabiate blossoms, a search was made for examples across the angiosperms and these were combined with personal observations collected during the last 25 years, coupled with knowledge from the literature. New functional terms are introduced that are independent of morphological and taxonomic associations.
Key Results
Bilabiate constructions occur in at least 38 angiosperm families. They are characterized by dorsiventral organization and dorsal pollen transfer. They are most often realised on the level of a single flower, but may also be present in an inflorescence or as part of a so-called ‘walk-around flower’. Interestingly, in functional terms all nototribic blossoms represent bilabiate constructions. The great majority of specialized bee-flowers can thus be included under bilabiate and keel blossoms. The syndrome introduced here, however, also paves the way for the inclusion of larger animals such as birds and bats. The most important evolutionary trends appear to be in the saving of pollen and the precision of its transfer. With special reference to the Lamiales, selected examples of bilabiate flowers are presented and their functional significance is discussed.
Bilabiate blossoms protect their pollen against pollen-collecting bees and at the same time render their pollination more precisely. The huge diversity of realised forms indicate the high selection pressure towards the bilabiate syndrome. As bees are very inventive, however, bilabiate constructions will not represent the ultimate response to bees.
PMCID: PMC2735325  PMID: 17652341
Bilabiate flowers; nototribic (dorsal) pollination; floral diversity; bee blossoms; functional morphology
20.  Why do so many petals have conical epidermal cells? 
Annals of Botany  2011;108(4):609-616.
The conical epidermal cells found on the petals of most Angiosperm species are so widespread that they have been used as markers of petal identity, but their function has only been analysed in recent years. This review brings together diverse data on the role of these cells in pollination biology.
The published effects of conical cells on petal colour, petal reflexing, scent production, petal wettability and pollinator grip on the flower surface are considered. Of these factors, pollinator grip has been shown to be of most significance in the well-studied Antirrhinum majus/bumble-bee system. Published data on the relationship between epidermal cell morphology and floral temperature were limited, so an analysis of the effects of cell shape on floral temperature in Antirrhinum is presented here. Statistically significant warming by conical cells was not detected, although insignificant trends towards faster warming at dawn were found, and it was also found that flat-celled flowers could be warmer on warm days. The warming observed is less significant than that achieved by varying pigment content. However, the possibility that the effect of conical cells on temperature might be biologically significant in certain specific instances such as marginal habitats or weather conditions cannot be ruled out.
Conical epidermal cells can influence a diverse set of petal properties. The fitness benefits they provide to plants are likely to vary with pollinator and habitat, and models are now required to understand how these different factors interact.
PMCID: PMC3170151  PMID: 21470973
Antirrhinum majus; conical cell; epidermis; floral scent; floral temperature; flower colour; grip; petal; pollination; wettability
21.  Domestication of cardamom (Elettaria cardamomum) in Western Ghats, India: divergence in productive traits and a shift in major pollinators 
Annals of Botany  2009;103(5):727-733.
Background and Aims
Elettaria cardamomum, a highly priced spice, is native to the Western Ghats of South India. Wild populations still occur in isolated patches in their natural habitats; however, much of today's commercial product comes from cultivated sources. There is no information on domestication-related traits of this species; the main objective of this study was to compare wild and cultivated populations of cardamom in terms of vegetative and reproductive features in order to identify domestication syndromes and to examine whether the two populations have developed reproductive barriers.
Two wild populations and five cultivated plantations were used for the present study. Vegetative and floral traits, flowering phenology, pollination biology and breeding systems of wild and cultivated populations were compared. Effective pollinators amongst floral visitors were identified by confirming pollen transfer as well as by fruit set following their visit to virgin flowers. Manual pollinations were carried out in order to study the breeding systems of the two populations and reproductive barriers, if any, between them.
Key Results
Several productive traits including the number of branches, number of inflorescences, and total number of flowers per clump, number of flowers that open each day, the duration of flowering, the length of the flower and the amount of nectar per flower are significantly greater in cultivated cardamom. The principal pollinators in wild cardamom are solitary bees, Megachile sp. and two species of Amegilla, whereas those in cultivated cardamom are the social bees Apis dorsata, A. cerana and Trigona iridipennis. Both the wild and cultivated populations are self-compatible and there are no reproductive barriers between the two populations.
Domestication in cardamom has brought about significant changes in vegetative and reproductive traits and a shift in effective pollinators from native solitary bees to social bees. The shift in pollinators seems to be due to the availability of a large number of flowers for prolonged periods in cultivated cardamom that can attract and sustain social bees, rather than due to co-evolution of the flower and the pollinator.
PMCID: PMC2707870  PMID: 19126635
Elettaria cardamomum; wild cardamom; domestication; Amegilla sp.; Apis cerana; Apis dorsata; Megachile sp.; pollination efficiency; solitary bees; social bees
22.  Geographical differentiation in floral traits across the distribution range of the Patagonian oil-secreting Calceolaria polyrhiza: do pollinators matter? 
Annals of Botany  2013;113(2):251-266.
Background and Aims
The underlying evolutionary processes of pollinator-driven floral diversification are still poorly understood. According to the Grant–Stebbins model speciation begins with adaptive local differentiation in the response to spatial heterogeneity in pollinators. Although this crucial process links the micro- and macroevolution of floral adaptation, it has received little attention. In this study geographical phenotypic variation was investigated in Patagonian Calceolaria polyrhiza and its pollinators, two oil-collecting bee species that differ in body size and geographical distribution.
Patterns of phenotypic variation were examined together with their relationships with pollinators and abiotic factors. Six floral and seven vegetative traits were measured in 45 populations distributed across the entire species range. Climatic and edaphic parameters were determined for 25 selected sites, 2–16 bees per site of the most frequent pollinator species were captured, and a critical flower–bee mechanical fitting trait involved in effective pollination was measured. Geographical patterns of phenotypic and environmental variation were examined using uni- and multivariate analyses. Decoupled geographical variation between corolla area and floral traits related to the mechanical fit of pollinators was explored using a Mantel test.
Key Results
The body length of pollinators and the floral traits related to mechanical fit were strongly correlated with each other. Geographical variation of the mechanical-fit-related traits was decoupled from variation in corolla size; the latter had a geographical pattern consistent with that of the vegetative traits and was mainly affected by climatic gradients.
The results are consistent with pollinators playing a key role in shaping floral phenotype at a geographical scale and promoting the differentiation of two floral ecotypes. The relationship between the critical floral-fit-related trait and bee length remained significant even in models that included various environmental variables and an allometric predictor (corolla area). The abiotic environment also has an important role, mainly affecting floral size. Decoupled geographical variation between floral mechanical-fit-related traits and floral size would represent a strategy to maintain plant–pollinator phenotypic matching in this environmentally heterogeneous area.
PMCID: PMC3890392  PMID: 24252281
Abiotic environmental gradients; bee morphology; Calceolaria; floral ecotypes; geographical range; local adaptation; oil-collecting bees; oil-offering flowers; Patagonia; phenotypic covariance; specialized pollination; speciation; vegetative morphology
23.  Bumblebees Learn Polarization Patterns 
Current Biology  2014;24(12):1415-1420.
Foraging insect pollinators such as bees must find and identify flowers in a complex visual environment. Bees use skylight polarization patterns for navigation [1–3], a capacity mediated by the polarization-sensitive dorsal rim area (DRA) of their eye [4, 5]. While other insects use polarization sensitivity to identify appropriate habitats [6], oviposition sites, and food sources [7], to date no nonnavigational functions of polarization vision have been identified in bees. Here we investigated the ability of bumblebees (Bombus terrestris) to learn polarization patterns on artificial “flowers” in order to obtain a food reward. We show that foraging bumblebees can learn to discriminate between two differently polarized targets, but only when the target artificial “flower” is viewed from below. A context for these results is provided by polarization imaging of bee-pollinated flowers, revealing the potential for polarization patterns in real flowers. Bees may therefore have the ability to use polarization vision, possibly mediated by their polarization-sensitive DRA, both for navigation and to learn polarization patterns on flowers, the latter being the first nonnavigational function for bee polarization vision to be identified.
•Bumblebees (Bombus terrestris) learn polarization patterns on artificial “flowers”•Polarization patterns were only learned from downward-facing, pendant “flowers”•Polarization vision in bumblebees is not restricted to sun-compass navigation•Polarization patterns of petals may be a component of floral signaling
Foraging bumblebees (Bombus terrestris) learn polarization patterns on downward-facing artificial “flowers” to find food, demonstrating that their polarization vision is not restricted to sun-compass navigation. Polarization patterns occur on the petals of real flowers and may be a, hitherto overlooked, component of floral signaling.
PMCID: PMC4062934  PMID: 24909321
24.  Stigma Development and Receptivity in Almond (Prunus dulcis) 
Annals of Botany  2006;97(1):57-63.
• Background and Aims Fertilization is essential in almond production, and pollination can be limiting in production areas. This study investigated stigma receptivity under defined developmental stages to clarify the relationship between stigma morphology, pollen germination, tube growth and fruit set.
• Methods Light and scanning electron microscopy were employed to examine stigma development at seven stages of flower development ranging from buds that were swollen to flowers in which petals were abscising. Flowers at different stages were hand pollinated and pollen germination and tube growth assessed. Artificial pollinations in the field were conducted to determine the effect of flower age on fruit set.
• Key Results Later stages of flower development exhibited greater stigma receptivity, i.e. higher percentages of pollen germination and more extensive tube growth occurred in older (those opened to the flat petal stage or exhibiting petal fall) than younger flowers. Enhanced stigma receptivity was associated with elongation of stigmatic papillae and increased amounts of stigmatic exudate that inundated papillae at later developmental stages. Field pollinations indicated that the stigma was still receptive and nut set was maintained in older flowers.
• Conclusions Stigma receptivity in almond does not become optimal until flowers are past the fully open stage. The stigma is still receptive and fruit set is maintained in flowers even at the stage when petals are abscising. Strategies to enhance pollination and crop yield, including the timing and placement of honey bees, should consider the effectiveness of developmentally advanced flowers.
PMCID: PMC2803376  PMID: 16287904
Almond; effective pollination period; Prunus dulcis; stigma receptivity; stigmatic exudate
25.  The North‐east‐Brazilian Liana, Adenocalymna dichilum (Bignoniaceae) Pollinated by Bats 
Annals of Botany  2004;93(5):609-613.
• Background and Aims Of the set of syndromes displayed by specialized (euphilic) flowers, adaptation to pollination by bats (chiropterophily) is the least known. Accumulated new evidence reveals that this pollination mode plays a considerable role in tropical communities, especially in the neotropics. One family in which bat‐pollinated species are known in several genera is the Bignoniaceae. Here is reported, for the first time, bat pollination and floral ecology in Adenocalymna dichilum (tribe Bignonieae).
• Methods Floral features of this species growing in Bahia (north‐east Brazil) indicated possible chiropterophily, which was subsequently confirmed by direct observation and from photographs of bat visits. Timing of anthesis and nectar parameters were monitored in the field, and floral morphology was investigated with fixed flowers.
• Key Results One to two flowers open per night on the upright, simple racemes of A. dichilum during several weeks in a ‘steady state’ mode. The bilabiate, cream‐coloured corollas are functional for only a single night and wilt during the following day. A stout corolla, with a musky odour, and a large nectary disc with large quantities of watery nectar also conform to the syndrome. Glossophaga soricina (Glossophaginae) visited and pollinated the flowers in a trap‐lining manner. Whilst hovering, the bats put their heads into the corolla mouth for less than 1 s to feed, thereby effecting the transfer of pollen which is deposited on their backs.
• Conclusions Adenocalymna, a New World genus comprising approx. 50 species, exhibits floral adaptive radiation including species pollinated by bees, birds and possibly moths. The discovery of chiropterophily in A. dichilum adds another facet to the array of floral syndromes represented in the genus.
PMCID: PMC4242311  PMID: 15037447
Adenocalymna; Bignoniaceae; chiropterophily; Glossophaga; Campo rupestre; north‐east Brazil

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