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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.
doi:10.1100/2012/381575
PMCID: PMC3361249  PMID: 22666114
2.  Reproductive success of non-rewarding Cypripedium japonicum benefits from low spatial dispersion pattern and asynchronous flowering 
Annals of Botany  2009;103(8):1227-1237.
Background and Aims
Outcrossing animal-pollinated plants, particularly non-rewarding species, often experience pollinator limitation to reproduction. Pollinator visitation is affected by various factors, and it is hypothesized that reproduction in non-rewarding plants would benefit from low spatial flower abundance and asynchronous flowering. In order to test this hypothesis, the influence of spatial pattern and flowering phenology on male and female reproductive success (RS) was investigated in a non-rewarding orchid, Cypripedium japonicum, in central China over two flowering seasons.
Methods
The probabilities of intrafloral self-pollination and geitonogamy caused by pollinator behaviours were estimated from field observations. Pollinator limitation was evaluated by hand-pollination experiments. RS was surveyed in different spatial flower dispersal patterns and local flower densities. The effects of flowering phenological traits on RS were assessed by univariate and multivariate regression analyses.
Key Results
Hand-pollination experiments revealed that fruit production was strongly pollen limited throughout the entire reproductive season – over two seasons, 74·3 % of individuals set fruit following hand pollination, but only 5·2–7·7 % did so under natural conditions. Intrafloral self-pollination and geitonogamy within the potential clones might be rare. Both male and female fitness were substantially lower in clustered plants than in those growing singly. An increase in local conspecific flower density significantly and negatively influenced male RS, but had no effect on female RS. Phenotypic selection analysis indicated that individuals flowering earlier have the greatest probability of RS. Over 85 % of sampled flowering individuals had a flowering synchrony value >0·7; however, highly synchronous flowering was not advantageous for RS, as indicated by the negative directional selection differentials and gradients, and by the positive quadratic selection gradients.
Conclusions
These results support the hypothesis that, as a consequence of density-dependent selection, low spatio-temporal flower abundance is advantageous for attracting pollinators and for reproduction in natural populations of non-rewarding C. japonicum.
doi:10.1093/aob/mcp066
PMCID: PMC2685325  PMID: 19318381
Cypripedium japonicum; pollen limitation; spatial dispersal pattern; flowering synchrony; reproductive success; selection differential; selection gradient; deceptive flower
3.  Reproductive isolation and pollination success of rewarding Galearis diantha and non-rewarding Ponerorchis chusua (Orchidaceae) 
Annals of Botany  2010;107(1):39-47.
Background and Aims
Increasing evidence challenges the conventional perception that orchids are the most distinct example of floral diversification due to floral or prezygotic isolation. Regarding the relationship between co-flowering plants, rewarding and non-rewarding orchids in particular, few studies have investigated whether non-rewarding plants affect the pollination success of rewarding plants. Here, floral isolation and mutual effects between the rewarding orchid Galearis diantha and the non-rewarding orchid Ponerorchis chusua were investigated.
Methods
Flowering phenological traits were monitored by noting the opening and wilting dates of the chosen individual plants. The pollinator pool and pollinator behaviour were assessed from field observations. Key morphological traits of the flowers and pollinators were measured directly in the field. Pollinator limitation and interspecific compatibility were evaluated by hand-pollination experiments. Fruit set was surveyed in monospecific and heterospecific plots.
Key Results
The species had overlapping peak flowering periods. Pollinators of both species displayed a certain degree of constancy in visiting each species, but they also visited other flowers before landing on the focal orchids. A substantial difference in spur size between the species resulted in the deposition of pollen on different regions of the body of the shared pollinator. Hand-pollination experiments revealed that fruit set was strongly pollinator-limited in both species. No significant difference in fruit set was found between monospecific plots and heterospecific plots.
Conclusions
A combination of mechanical isolation and incomplete ethological isolation eliminates the possibility of pollen transfer between the species. These results do not support either the facilitation or competition hypothesis regarding the effect of nearby rewarding flowers on non-rewarding plants. The absence of a significant effect of non-rewarding P. chusua on rewarding G. diantha can be ascribed to low levels of overlap between the pollinator pools of two species.
doi:10.1093/aob/mcq210
PMCID: PMC3002470  PMID: 20961923
Galearis diantha; Ponerorchis chusua; rewarding and non-rewarding; mechanical isolation; ethological isolation; pollinator limitation; fruit set
4.  Plant Pollinator Networks along a Gradient of Urbanisation 
PLoS ONE  2013;8(5):e63421.
Background
Habitat loss is one of the principal causes of the current pollinator decline. With agricultural intensification, increasing urbanisation is among the main drivers of habitat loss. Consequently studies focusing on pollinator community structure along urbanisation gradients have increased in recent years. However, few studies have investigated how urbanisation affects plant-pollinator interaction networks. Here we assessed modifications of plant-pollinator interactions along an urbanisation gradient based on the study of their morphological relationships.
Methodology/Principal Findings
Along an urbanisation gradient comprising four types of landscape contexts (semi-natural, agricultural, suburban, urban), we set up experimental plant communities containing two plant functional groups differing in their morphological traits (“open flowers” and “tubular flowers”). Insect visitations on these communities were recorded to build plant-pollinator networks. A total of 17 857 interactions were recorded between experimental plant communities and flower-visitors. The number of interactions performed by flower-visitors was significantly lower in urban landscape context than in semi-natural and agricultural ones. In particular, insects such as Syrphidae and solitary bees that mostly visited the open flower functional group were significantly impacted by urbanisation, which was not the case for bumblebees. Urbanisation also impacted the generalism of flower-visitors and we detected higher interaction evenness in urban landscape context than in agricultural and suburban ones. Finally, in urban context, these modifications lowered the potential reproductive success of the open flowers functional group.
Conclusions/Significance
Our findings show that open flower plant species and their specific flower-visitors are especially sensitive to increasing urbanisation. These results provide new clues to improve conservation measures within urbanised areas in favour of specialist flower-visitors. To complete this functional approach, studies using networks resolved to the species level along urbanised gradients would be required.
doi:10.1371/journal.pone.0063421
PMCID: PMC3661593  PMID: 23717421
5.  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.
doi:10.1371/journal.pone.0093615
PMCID: PMC3972141  PMID: 24690875
6.  Ability of Bumblebees to Discriminate Differences in the Shape of Artificial Flowers of Primula sieboldii (Primulaceae) 
Annals of Botany  2007;99(6):1175-1182.
Background and Aims
Flower shapes are important visual cues for pollinators. However, the ability of pollinators to discriminate between flower shapes under natural conditions is poorly understood. This study focused on the diversity of flower shape in Primula sieboldii and investigated the ability of bumblebees to discriminate between flowers by combining computer graphics with a traditional behavioural experiment.
Methods
Elliptic Fourier descriptors described shapes by transforming coordinate information for the contours into coefficients, and principal components analysis summarized these coefficients. Using these methods, artificial flowers were created based on the natural diversity of petal shape in P. sieboldii. Dual-choice tests were then performed to investigate the ability of the bumblebees to detect differences in the aspect ratio of petals and the depth of their head notch.
Key Results
The insects showed no significant ability to detect differences in the aspect ratio of the petals under natural conditions unless the morphological distance increased to an unrealistic level. These results suggest the existence of a perception threshold for distances in this parameter. The bumblebees showed a significant preference for narrow petals even after training using flowers with wide petals. The bumblebees showed a significant ability to discriminate based on the depth of the petal head notch after training using artificial flowers with a deep head notch. However, they showed no discrimination in tests with training using extreme distances between flowers in this parameter.
Conclusions
A new type of behavioural experiment was demonstrated using real variation in flower corolla shape in P. sieboldii. If the range in aspect ratios of petals expands much further, bumblebees may learn to exhibit selective behaviour. However, because discrimination by bumblebees under natural conditions was low, there may be no strong selective behaviour based on innate or learned preferences under natural conditions.
doi:10.1093/aob/mcm059
PMCID: PMC3243577  PMID: 17553825
Bumblebee; computer graphics; elliptic Fourier descriptors; flower corolla shape; Primula sieboldii; principal components analysis; visual cue
7.  Functional Androdioecy in Critically Endangered Gymnocladus assamicus (Leguminosae) in the Eastern Himalayan Region of Northeast India 
PLoS ONE  2014;9(2):e87287.
Gymnocladus assamicus is a critically endangered tree species endemic to Northeast India, and shows sexual dimorphism with male and hermaphrodite flowers on separate trees. We studied phenology, reproductive biology and mating system of the species. The flowers are small, tubular, odorless and last for about 96 hours. Pollen grains in both morphs were viable and capable of fertilization leading to fruit and seed set. Scanning electron micrographs revealed morphologically similar pollen in both male and hermaphrodite flowers. The fruit set in open pollinated flowers was 43.61 percent, while controlled autogamous and geitonogamous pollinations yielded 76.81 and 65.58 percent fruit set respectively. Xenogamous pollinations between male and hermaphrodite flowers resulted in 56.85 percent fruit set and pollinations between hermaphrodite flowers yielded 67.90 percent fruit set. This indicates a functionally androdioecious mating system and pollination limited fruit set in G. assamicus. Phylogenetic analyses of Gymnocladus and the sister genus Gleditsia are needed to assess if the androdioecious mating system in G. assamicus evolved from dioecy as a result of selection for hermaphrodites for reproductive assurance during colonization of pollination limited high altitude ecosystems.
doi:10.1371/journal.pone.0087287
PMCID: PMC3938411  PMID: 24586267
8.  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.
doi:10.1093/aob/mcj013
PMCID: PMC2803376  PMID: 16287904
Almond; effective pollination period; Prunus dulcis; stigma receptivity; stigmatic exudate
9.  Comparative Anatomy and Morphology of Nectar-producing Melastomataceae 
Annals of Botany  2008;102(6):899-909.
Background and Aims
Most neotropical Melastomataceae have bee-pollinated flowers with poricidal anthers. However, nectar rewards are known to be produced in about 80 species in eight genera from four different tribes. These nectar-producing species are pollinated by both vertebrates and invertebrates.
Methods
The floral morphology and anatomy of 14 species was studied in six genera of nectar-producing Melastomataceae (Blakea, Brachyotum, Charianthus, Huilaea, Meriania and Miconia). Anatomical methods included scanning electron microscopy, and serial sections of paraffin-embedded flowers.
Key Results
All vertebrate-pollinated melastome flowers have petals that do not open completely at anthesis, thus forming a pseudo-tubular corolla, while closely related species that are bee pollinated have rotate or reflexed corollas. In most species, nectar secretion is related to stomatal or epidermal nectaries and not filament slits as previously reported. Moreover, the nectar is probably supplied by large vascular bundles near the release area. Blakea and Huilaea have nectary stomata located upon the dorsal anther connective appendages. Brachyotum also has nectary stomata on the anther connectives, but these are distributed lengthwise along most of the connective. Meriania may release nectar through the anther connective, but has additional nectary stomata on the inner walls of the hypanthium. Miconia has nectary stomata on the ovary apex. Charianthus nectaries were not found, but there is circumstantial evidence that nectar release occurs through the epidermis at the apex of the ovary and the lower portions of the inner wall of the hypanthium.
Conclusions
Nectar release in Melastomataceae is apparently related to nectary stomata and not filament slits. The presence of nectary stomata on stamens and on ovary apices in different lineages suggests that the acquisition of nectaries is a derived condition. Nectary location also supports a derived condition, because location is strongly consistent within each genus, but differs between genera.
doi:10.1093/aob/mcn180
PMCID: PMC2712398  PMID: 18819949
Blakea; Brachyotum; Charianthus; Huilaea; Meriania; Melastomataceae; Miconia; nectaries; nectary stomata; pollination
10.  Evolution of petal epidermal micromorphology in Leguminosae and its use as a marker of petal identity 
Annals of Botany  2009;104(6):1099-1110.
Background and Aims
The legume flower is highly variable in symmetry and differentiation of petal types. Most papilionoid flowers are zygomorphic with three types of petals: one dorsal, two lateral and two ventral petals. Mimosoids have radial flowers with reduced petals while caesalpinioids display a range from strongly zygomorphic to nearly radial symmetry. The aims are to characterize the petal micromorphology relative to flower morphology and evolution within the family and assess its use as a marker of petal identity (whether dorsal, lateral or ventral) as determined by the expression of developmental genes.
Methods
Petals were analysed using the scanning electron microscope and light microscope. A total of 175 species were studied representing 26 tribes and 89 genera in all three subfamilies of the Leguminosae.
Key Results
The papilionoids have the highest degree of variation of epidermal types along the dorsiventral axis within the flower. In Loteae and genistoids, in particular, it is common for each petal type to have a different major epidermal micromorphology. Papillose conical cells are mainly found on dorsal and lateral petals. Tabular rugose cells are mainly found on lateral petals and tabular flat cells are found only in ventral petals. Caesalpinioids lack strong micromorphological variation along this axis and usually have only a single major epidermal type within a flower, although the type maybe either tabular rugose cells, papillose conical cells or papillose knobby rugose cells, depending on the species.
Conclusions
Strong micromorphological variation between different petals in the flower is exclusive to the subfamily Papilionoideae. Both major and minor epidermal types can be used as micromorphological markers of petal identity, at least in papilionoids, and they are important characters of flower evolution in the whole family. The molecular developmental pathway between specific epidermal micromorphology and the expression of petal identity genes has yet to be established.
doi:10.1093/aob/mcp211
PMCID: PMC2766202  PMID: 19789174
Epidermis; Fabaceae; Papilionoideae; Caesalpinioideae; Mimosoideae; petal surface; scanning electron microscopy; papillose conical cells; tabular rugose cells; tabular flat cells; organ identity
11.  Sterile flowers increase pollinator attraction and promote female success in the Mediterranean herb Leopoldia comosa 
Annals of Botany  2012;111(1):103-111.
Background and Aims
Large floral displays have opposing consequences for animal-pollinated angiosperms: they attract more pollinators but also enable elevated among-flower self-pollination (geitonogamy). The presence of sterile flowers as pollinator signals may enhance attraction while allowing displays of fewer open fertile flowers, limiting geitonogamy. The simultaneous contributions of fertile and non-fertile display components to pollinator attraction and reproductive output remain undetermined.
Methods
The simultaneous effects of the presence of sterile flowers and fertile-flower display size in two populations of Leopoldia comosa were experimentally assessed. Pollinator behaviour, pollen removal and deposition, and fruit and seed production were compared between intact plants and plants with sterile flowers removed.
Key Results
The presence of sterile flowers almost tripled pollinator attraction, supplementing the positive effect of the number of fertile flowers on the number of bees approaching inflorescences. Although attracted bees visited more flowers on larger inflorescences, the number visited did not additionally depend on the presence of sterile flowers. The presence of sterile flowers improved all aspects of plant performance, the magnitude of plant benefit being context dependent. During weather favourable to pollinators, the presence of sterile flowers increased pollen deposition on stigmas of young flowers, but this difference was not evident in older flowers, probably because of autonomous self-pollination in poorly visited flowers. Total pollen receipt per stigma decreased with increasing fertile display size. In the population with more pollinators, the presence of sterile flowers increased fruit number but not seed set or mass, whereas in the other population sterile flowers enhanced seeds per fruit, but not fruit production. These contrasts are consistent with dissimilar cross-pollination and autonomous self-pollination, coupled with the strong predispersal inbreeding depression exhibited by L. comosa populations.
Conclusions
Sterile flowers enrich pollination quality by promoting pollen export and import, while limiting the mating costs of geitonogamy associated with large fertile displays.
doi:10.1093/aob/mcs243
PMCID: PMC3523656  PMID: 23131298
Anthophora; cross-pollination; geitonogamy; fertile floral display; mating cost; Leopoldia comosa; non-fertile flowers; outcrossing; pollen deposition; pollen quality; pollen removal; sterile flowers
12.  Differences in dichogamy and herkogamy contribute to higher selfing in contrasting environments in the annual Blackstonia perfoliata (Gentianaceae) 
Annals of Botany  2013;111(4):651-661.
Background and Aims
The establishment of plant populations in novel environments may generate pronounced shifts in floral traits and plant mating systems, particularly when pollinators are scarce. In this study, floral morphology and mating system functioning are compared between recently established and older populations of the annual plant Blackstonia perfoliata that occur in different pollinator environments.
Methods
Hand-pollination and emasculation experiments were conducted to assess the extent of pollinator-mediated pollen deposition and pollen limitation, and the contribution of autonomous selfing to total seed production. Detailed measurements of key floral traits were performed to compare the flower morphology and mating system functioning between plants from both pollination environments.
Key Results
Pollinator-mediated pollen deposition was about twice as low in the recently colonized and pollinator-poor environment compared with the old and pollinator-rich sites, but total seed set was little affected by any type of pollen limitation. The contribution of autonomous selfing to total seed production was higher in the pollinator-poor sites than in the pollinator-rich sites (index of reproductive assurance = 0·56 and 0·17, respectively), and seed production was only poorly affected by selfing, whereas in the pollinator-rich populations selfing reduced total reproductive output by about 40 % compared with outcross pollination. Plants originating from pollinator-poor environments produced smaller flowers that showed significantly lower levels of dichogamy (i.e. protogyny) and herkogamy. These reductions resulted in a 2-fold higher capacity for autonomous selfing under pollinator-free conditions (index of autonomous selfing = 0·81 and 0·41 in plants originating from the pollinator-poor and pollinator-rich environment, respectively).
Conclusions
The results illustrate that plant populations colonizing novel environments can differ markedly in floral morphology and mating system functioning. Due to a temporal shift in the male phase, the breeding system of B. perfoliata shifted from delayed selfing under pollinator-rich conditions towards competing selfing in recently established populations, providing greater reproductive assurance when pollinators and/or reproductive partners are limited.
doi:10.1093/aob/mct031
PMCID: PMC3605963  PMID: 23408833
Autogamy; competing selfing; delayed selfing; dichogamy; herkogamy; pollen limitation; pollen quality; reproductive assurance; Blackstonia perfoliata; Gentianaceae
13.  Spatial variation in selection on corolla shape in a generalist plant is promoted by the preference patterns of its local pollinators 
Summary
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.
doi:10.1098/rspb.2008.0512
PMCID: PMC2603243  PMID: 18544510
corolla shape evolution; pollinator preference; spatial variation; geometric morphometrics; nectar; pollen
14.  The role of short-tongued insects in floral variation across the range of a style-dimorphic plant 
Annals of Botany  2012;111(2):317-328.
Background and Aims
Heterostyly and related style polymorphisms are suitable model systems to evaluate the importance of functional pollinators in the maintenance of population variability. In Narcissus papyraceus different functional pollinators, incompatibility system and flower morphology have been proposed to influence the maintenance of polymorphism through their effect on disassortative mating. Here a test is done to find out if the visitation rate of long- versus short-tongued pollinators correlates with the morph ratio and if the latter is related to other flower traits of the species across its main geographic range.
Methods
Floral traits from 34 populations in the south-west of the Iberian Peninsula and in north-west Africa were measured, perianth variation was described and a comparison was made of allometric relationships between sex organs and floral tube. Correlations between pollinator guilds, stigma–anther separation of reciprocal morphs (our proxy for disassortative mating) and morph-ratio variation were analysed. Finally, the incompatibility system of the species in the northern and southern borders of its distribution are described.
Key Results
Flowers from southern populations were significantly larger than flowers from centre and northern populations. The abundance of short-styled plants decreased gradually with increasing distance from the core region (the Strait of Gibraltar), with these disappearing only in the northern range. Although there was a significant difference in stigma–anther separation among populations, morph ratio was not associated with reciprocity or floral tube length. Long-style morph frequency increased with short-tongued pollinator visitation rate. Populations from both edges of the distribution range were self-incompatible and within- and between-morph compatible.
Conclusions
The style morph ratio changed gradually, whereas perianth trait variation showed abrupt changes with two morphotypes across the range. The positive relationship between the visitation rate of short-tongued pollinators and the decrease of the short-style morph supports our initial hypothesis. The results highlight the importance of different pollinators in determining the presence of style polymorphism.
doi:10.1093/aob/mcs261
PMCID: PMC3555527  PMID: 23223205
Disassortative mating; floral morphology; geographic variation; morph ratio; Narcissus papyraceus; pollinators; stylar dimorphism
15.  A global test of the pollination syndrome hypothesis 
Annals of Botany  2009;103(9):1471-1480.
Background and Aims
‘Pollination syndromes’ are suites of phenotypic traits hypothesized to reflect convergent adaptations of flowers for pollination by specific types of animals. They were first developed in the 1870s and honed during the mid 20th Century. In spite of this long history and their central role in organizing research on plant–pollinator interactions, the pollination syndromes have rarely been subjected to test. The syndromes were tested here by asking whether they successfully capture patterns of covariance of floral traits and predict the most common pollinators of flowers.
Methods
Flowers in six communities from three continents were scored for expression of floral traits used in published descriptions of the pollination syndromes, and simultaneously the pollinators of as many species as possible were characterized.
Key Results
Ordination of flowers in a multivariate ‘phenotype space’ defined by the syndromes showed that almost no plant species fall within the discrete syndrome clusters. Furthermore, in approximately two-thirds of plant species, the most common pollinator could not be successfully predicted by assuming that each plant species belongs to the syndrome closest to it in phenotype space.
Conclusions
The pollination syndrome hypothesis as usually articulated does not successfully describe the diversity of floral phenotypes or predict the pollinators of most plant species. Caution is suggested when using pollination syndromes for organizing floral diversity, or for inferring agents of floral adaptation. A fresh look at how traits of flowers and pollinators relate to visitation and pollen transfer is recommended, in order to determine whether axes can be identified that describe floral functional diversity more successfully than the traditional syndromes.
doi:10.1093/aob/mcp031
PMCID: PMC2701765  PMID: 19218577
Convergent evolution; floral traits; global; montane meadow; multidimensional scaling; mutualism; phenotype space; pollination syndromes; temperate grassland; test; tropical forest; tropical mountains
16.  Why do so many petals have conical epidermal cells? 
Annals of Botany  2011;108(4):609-616.
Background
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.
Scope
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.
Conclusions
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.
doi:10.1093/aob/mcr065
PMCID: PMC3170151  PMID: 21470973
Antirrhinum majus; conical cell; epidermis; floral scent; floral temperature; flower colour; grip; petal; pollination; wettability
17.  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.
Methods
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.
Conclusions
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.
doi:10.1093/aob/mcr268
PMCID: PMC3241588  PMID: 22025522
Morrenia odorata; Morrenia brachystephana; asclepiads; functional morphology; pollination mechanism; wasps; pollinator effectiveness
18.  Orthoptera, a new order of pollinator 
Annals of Botany  2010;105(3):355-364.
Background and Aims
Pollinator-mediated selection and evolution of floral traits have long fascinated evolutionary ecologists. No other plant family shows as wide a range of pollinator-linked floral forms as Orchidaceae. In spite of the large size of this model family and a long history of orchid pollination biology, the identity and specificity of most orchid pollinators remains inadequately studied, especially in the tropics where the family has undergone extensive diversification. Angraecum (Vandeae, Epidendroideae), a large genus of tropical Old World orchids renowned for their floral morphology specialized for hawkmoth pollination, has been a model system since the time of Darwin.
Methods
The pollination biology of A. cadetii, an endemic species of the islands of Mauritius and Reunion (Mascarene Islands, Indian Ocean) displaying atypical flowers for the genus (white and medium-size, but short-spurred) was investigated. Natural pollinators were observed by means of hard-disk camcorders. Pollinator-linked floral traits, namely spur length, nectar volume and concentration and scent production were also investigated. Pollinator efficiency (pollen removal and deposition) and reproductive success (fruit set) were quantified in natural field conditions weekly during the 2003, 2004 and 2005 flowering seasons (January to March).
Key Results
Angraecum cadetii is self-compatible but requires a pollinator to achieve fruit set. Only one pollinator species was observed, an undescribed species of raspy cricket (Gryllacrididae, Orthoptera). These crickets, which are nocturnal foragers, reached flowers by climbing up leaves of the orchid or jumping across from neighbouring plants and probed the most ‘fresh-looking’ flowers on each plant. Visits to flowers were relatively long (if compared with the behaviour of birds or hawkmoths), averaging 16·5 s with a maximum of 41·0 s. At the study site of La Plaine des Palmistes (Pandanus forest), 46·5 % of flowers had pollen removed and 27·5 % had pollinia deposited on stigmas. The proportion of flowers that set fruit ranged from 11·9 % to 43·4 %, depending of the sites sampled across the island.
Conclusions
Although orthopterans are well known for herbivory, this represents the first clearly supported case of orthopteran-mediated pollination in flowering plants.
doi:10.1093/aob/mcp299
PMCID: PMC2826249  PMID: 20067913
Angraecum; Mascarene Archipelago; oceanic islands; Orchidaceae; Orthoptera; plant–pollinator interactions; pollinator shifts
19.  Comparative evolution of flower and fruit morphology 
Angiosperm diversification has resulted in a vast array of plant morphologies. Only recently has it been appreciated that diversification might have proceeded quite differently for the two key diagnostic structures of this clade, flowers and fruits. These structures are hypothesized to have experienced different selective pressures via their interactions with animals in dispersal mutualisms, resulting in a greater amount of morphological diversification in animal-pollinated flowers than in animal-dispersed fruits. I tested this idea using size and colour traits for the flowers and fruits of 472 species occurring in three floras (St John, Hawaii and the Great Plains). Phylogenetically controlled analyses of nearest-neighbour distances in multidimensional trait space matched the predicted pattern: in each of the three floras, flowers were more divergent from one another than were fruits. In addition, the spacing of species clusters differed for flowers versus fruits in the flora of St John, with clusters in flower space more divergent than those in fruit space. The results are consistent with the idea that a major driver of angiosperm diversification has been stronger selection for divergent floral morphology than for divergent fruit morphology, although genetic, physiological and ecological constraints may also play a role.
doi:10.1098/rspb.2009.0483
PMCID: PMC2817210  PMID: 19474045
angiosperm radiation; floral character displacement; ecological sorting; pollinator constancy; frugivory; seed dispersal and pollination syndromes
20.  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.
Methods
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.
Conclusions
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.
doi:10.1093/aob/mcr077
PMCID: PMC3101143  PMID: 21498566
Vaccinium corymbosum; honey bees; bumble bees; volatile organic compounds; diurnal rhythm; nectar production; site of emission
21.  Visibility vs. biomass in flowers: exploring corolla allocation in Mediterranean entomophilous plants 
Annals of Botany  2009;103(7):1119-1127.
Background and Aims
While pollinators may in general select for large, morphologically uniform floral phenotypes, drought stress has been proposed as a destabilizing force that may favour small flowers and/or promote floral variation within species.
Methods
The general validity of this concept was checked by surveying a taxonomically diverse array of 38 insect-pollinated Mediterranean species. The interplay between fresh biomass investment, linear size and percentage corolla allocation was studied. Allometric relationships between traits were investigated by reduced major-axis regression, and qualitative correlates of floral variation explored using general linear-model MANOVA.
Key Results
Across species, flowers were perfectly isometrical with regard to corolla allocation (i.e. larger flowers were just scaled-up versions of smaller ones and vice versa). In contrast, linear size and biomass varied allometrically (i.e. there were shape variations, in addition to variations in size). Most floral variables correlated positively and significantly across species, except corolla allocation, which was largely determined by family membership and floral symmetry. On average, species with bilateral flowers allocated more to the corolla than those with radial flowers. Plant life-form was immaterial to all of the studied traits. Flower linear size variation was in general low among conspecifics (coefficients of variation around 10 %), whereas biomass was in general less uniform (e.g. 200–400 mg in Cistus salvifolius). Significant among-population differences were detected for all major quantitative floral traits.
Conclusions
Flower miniaturization can allow an improved use of reproductive resources under prevailingly stressful conditions. The hypothesis that flower size reflects a compromise between pollinator attraction, water requirements and allometric constraints among floral parts is discussed.
doi:10.1093/aob/mcp046
PMCID: PMC2707908  PMID: 19258340
Allometry; biomass; corolla; drought; evolution; flower; Mediterranean; sclerophyllous; size; variation; water
22.  Morphological and molecular identification to secure cultivar maintenance and management of self-sterile Rubus arcticus 
Annals of Botany  2013;111(4):713-721.
Background and Aims
Preservation of cultivar purity creates a particular challenge for plants that are self-incompatible, require insects for cross-pollination, and have easily germinating seeds and vigorously spreading rhizomes. As the fields must be planted with mixed populations, and a balance must be maintained between the cultivars to achieve effective pollination, methods for field monitoring of the relative density of different cultivars must be practical. Furthermore, a DNA-based method is needed for cultivar verification in the collections and outside of the growing season. The aim of this study was to develop both types of methods for Rubus arcticus (arctic bramble).
Methods
Morphological parameters were measured from six cultivars grown on three farms. Observations from the flowers and fruits included: petal and sepal number, flower diameter, arrangement of petals, size of calyx in relation to corolla, fruit weight, yield and soluble sugars. Observations from the leaves included: width and height of middle leaflet, shape of the base of terminal leaflet, shape of terminal leaflet, leaf margin serration and fingertip touch. The applicability of simple sequence repeat (SSR) or microsatellite DNA markers developed for red raspberry was tested on eight arctic bramble cultivars.
Key Results and Conclusions
Morphological and molecular identification methods were developed for R. arcticus. The best morphological characteristics were the length-to-width ratio of the middle leaflet and leaf margin serration. A particular characteristic, fingertip touch, was shown by electron microscopy to be related to the density and quality of the leaf hairs. Red raspberry SSR marker no. 126 proved to be applicable for differentiation of the eight arctic bramble cultivars tested. These identification methods are critical to secure the maintenance and management of R. arcticus. However, the challenges faced and approaches taken are equally applicable to other species with similar biology.
doi:10.1093/aob/mct029
PMCID: PMC3605961  PMID: 23456688
Arctic bramble; authenticity; cultivar purity; identification; morphological markers; Rubus arcticus ssp. arcticus; self-incompatibility; SSR markers; UPOV; preservation; cultivar maintenance; genetic resources conservation
23.  The Response of Spartium junceum Roots to Slope: Anchorage and Gene Factors 
Annals of Botany  2006;97(5):857-866.
• Background and Aims Plant anchorage is governed by complex, finely regulated mechanisms that occur at a morphological, architectural and anatomical level. Spanish broom (Spartium junceum) is a woody plant frequently found on slopes—a condition that affects plant anchorage. This plant grows throughout the Mediterranean area where it plays an important role in preventing landslides. Spanish broom seedlings respond promptly to slope by altering stem and root morphology. The aim of this study was to investigate the mechanisms whereby the root system of Spanish broom seedlings adapts to ensure anchorage to the ground.
• Methods Seedlings were grown in tilted and untilted pots under controlled conditions. The root apparatus was removed at different times of growth and subjected to morphological, biomechanical and molecular analyses.
• Key Results In slope-grown seedlings, changes in root system morphology, pulling strength and chemical lignin content, all features related to plant anchorage in the soil, were related to seedling age. cDNA-AFLP analysis revealed changes in the expression of several genes in root systems of slope-grown plants. BLAST analysis showed that some differentially expressed genes are homologues of genes induced by environmental stresses in other plant species, and/or are involved in the production of strengthening materials.
• Conclusion Plants use various mechanisms/strategies to respond to slope depending on their developmental stage.
doi:10.1093/aob/mcj603
PMCID: PMC2803426  PMID: 16352708
Anchorage; cDNA-AFLP; lignin; pulling; root morphology; root; slope; Spartium junceum
24.  Modification of flower architecture during early stages in the evolution of self-fertilization 
Annals of Botany  2009;103(6):951-962.
Background and Aims
The evolution of selfing from outcrossing is characterized by a series of morphological changes to flowers culminating in the selfing syndrome. However, which morphological traits initiate increased self-pollination and which are accumulated after self-fertilization establishes is poorly understood. Because the expression of floral traits may depend on the conditions experienced by an individual during flower development, investigation of changes in mating system should also account for environmental and developmental factors. Here, early stages in the evolution of self-pollination are investigated by comparing floral traits among Brazilian populations of Eichhornia paniculata (Pontederiaceae), an annual aquatic that displays variation in selfing rates associated with the breakdown of tristyly to semi-homostyly.
Methods
Thirty-one Brazilian populations under uniform glasshouse conditions were compared to investigate genetic and environmental influences on flower size and stigma–anther separation (herkogamy), two traits that commonly vary in association with transitions to selfing. Within-plant variation in herkogamy was also examined and plants grown under contrasting environmental conditions were compared to examine to what extent this trait exhibits phenotypic plasticity.
Key Results
In E. paniculata a reduction in herkogamy is the principal modification initiating the evolution of selfing. Significantly, reduced herkogamy was restricted to the mid-styled morph and occurred independently of flower size. Significant genetic variation for herkogamy was detected among populations and families, including genotypes exhibiting developmental instability of stamen position with bimodal distributions of herkogamy values. Cloned genets exposed to contrasting growth conditions demonstrated environmental control of herkogamy and genotypic differences in plasticity of this trait.
Conclusions
The ability to modify herkogamy independently of other floral traits, genetic variation in the environmental sensitivity of herkogamy, and the production of modified and unmodified flowers within some individuals, reveal the potential for dynamic control of the mating system in a species that commonly confronts heterogeneous aquatic environments.
doi:10.1093/aob/mcp015
PMCID: PMC2707899  PMID: 19202135
Eichhornia paniculata; expressivity; flower morphology; herkogamy; phenotypic plasticity; pleiotropy; population variation; self-fertilization; stigma–anther separation; outcrossing; tristyly
25.  Pollinator-mediated natural selection in Penstemon digitalis 
Plant Signaling & Behavior  2010;5(12):1688-1690.
Measuring the agents of natural selection is important because it allows us to understand not only which traits are expected to evolve but also why they will evolve. Natural selection by pollinators on floral traits is often assumed because in outcrossing animal-pollinated plants flowers are generally thought to function as advertisements of rewards directed at pollinators. We tested the role of bee pollinators in selection on Penstemon digitalis and found that pollinators were driving selection for larger and more flowers. However, what makes our publication unique is the additional information we gained from reviewing the few other studies that also directly tested whether pollinators were agents of selection on floral traits. As we would expect if pollinators are important agents of selection, selection on floral traits was significantly stronger when pollinators were present than when their choices were experimentally removed. Taken together, these results suggest that pollinators can be important drivers of selection in contemporary populations.
doi:10.4161/psb.5.12.14113
PMCID: PMC3115138  PMID: 21150291
plant vigor; Penstemon; pollinators; natural selection; selection gradient

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