Variation of nectar production in relation to plant characteristics in protandrous Aconitum gymnandrum

Aims Floral nectar plays a vital role in plant reproductive success by attracting pollinators. Nectar traits of a flower can depend directly on plant characteristics other than environmental factors and exhibit extensive flower- and plant-level variations. Studies on nectar traits frequently focused on intraplant variation for dichogamous plants, but few have paid attention to both intra- and interplant nectar variations in relation to plant characteristics. Revealing within- and among-plant variation and its relative magnitude is important for our understanding of how pollinator-mediated selection can act on nectar traits and evolution of nectar traits.Methods Through investigating protandrous Aconitum gymnandrum populations at the Alpine Meadows and Wetland Ecosystems Research Station of Lanzhou University, we examined the relationships between nectar production per flower and plant characteristics (e.g. flower position within inflorescences, floral sexual phases, flowering time, inflorescence size and floral attractive traits).Important findings A. gymnandrum exhibited a declining gradient in the nectar volume along inflorescences, with more nectar in basal flowers than distal ones. Protandrous flowers of A. gymnandrum did not show gender-biased nectar production while the nectar volume varied with different stages of floral sexual phases. The significant correlation between the first flowering date of individuals and the mean nectar volume per flower was positive in 2013, but became negative in 2014, suggesting complex effects of biotic and abiotic factors. The mean nectar volume per flower was not related to inflorescence size (the number of total flowers per plant). Furthermore, nectar production was weakly associated with floral attractive traits (the petal width and the galea height), even if the effect of flowering time of individuals was removed, suggesting that the honesty of floral traits as signals of nectar reward for pollinators is not stable in this species.     

The influence of plant spacing on density-dependent versus frequency-dependent spore transmission of the anther smut Microbotryum violaceum

The anther smut fungus Microbotryum violaceum is a pollinator-transmitted plant disease. As for other vector-borne diseases, frequency-dependent transmission patterns are predicted, in contrast to the density-dependent transmission of passively spread diseases. Frequency dependence will, however, only arise if vectors compensate for varying plant spacings. To test this assumption, we set up experimental populations of the host plant, Silene latifolia, with varying disease density (number of diseased plants per plot) and frequency (proportion of plants diseased), and three different plant spacings. We measured spore deposition on healthy flowers in these plots on two dates. Spore deposition decreased considerably from the first to the second census, perhaps related to the concomitant decrease in inflorescence sizes of diseased plants. At our first census, spore deposition rates varied with disease frequency, and the effect of frequency depended on plant spacing. While spore deposition was positively frequency dependent at the 1.5-m inter-plant spacing, no effect of disease frequency was found at a spacings of 0.5 m or 3 m. Nor was there an effect of disease density on spore deposition at the first census. At the later census, on the other hand, spore deposition increased almost significantly with increasing disease density (P = 0.08). This difference in deposition pattern together with a significant decrease in spore receipt indicates changes in pollinator spectrum and/or activity. The correlation of spore numbers among flowers within plants, an indication for intra-plant moves by vectors, was significant at 0.5 m and 1.5 m but not at 3 m. Floral traits and sex of individual plants influenced the number of spores they received. On the first census date, spore deposition increased with increasing inflorescence size in female but not in male plants. On the second census date, neither sex nor number of open flowers had an effect on spore receipt. None of the experimental plants became infected, however, probably because of the unusually hot and dry weather. Received: 14 May 1998 / Accepted: 19 November 1998     

Selection for floral integration and trait variation in zygomorphic flowers of Aconitum japonicum ssp. subcuneatum (Ranunculaceae)

Pollinator-mediated selection might lead to among-trait differences in the degree and pattern of floral integration and intra-flower variation. To examine the patterns of intra-flower variation in floral traits, including nectar volume, we performed a field study using the zygomorphic flowers of Aconitum japonicum ssp. subcuneatum. We investigated (1) correlations between the sizes of the left and right sepals and petals, (2) variation in floral traits among plants, within plants and within flowers, (3) effects of sexual phases on floral integration variation in floral and nectar traits, and (4) the effect of size and intra-flower variation in traits of the left and right sepals and petals on pollen removal by pollinators. Lateral sepal area, but not lower sepal area, was highly correlated between the left and right sepals. Floral traits were more integrated during the male phase than during the female phase. Nectar standing crop in male-phase flowers correlated with helmet height and lateral and lower sepal area, but in female-phase flowers it only correlated with spur length. While intra-flower variance in lateral sepal area accounted for approximately 10% of the overall variance in these traits, the variance in lower sepal area accounted for 70% of the overall variance. Lateral sepal area had a negative effect on the number of pollen grains remaining after pollinator visits. Low variance in lateral sepals within flowers and measurements of pollen removal suggest that lateral sepals play a more important role in pollen export than the other traits. Left and right sepals may be the targets of selection for symmetry in zygomorphic flowers.

     

Strong pollinator‐mediated selection for increased flower brightness and contrast in a deceptive orchid

Contrasting flower color patterns that putatively attract or direct pollinators toward a reward are common among angiosperms. In the deceptive orchid Anacamptis morio, the lower petal, which makes up most of the floral display, has a light central patch with dark markings. Within populations, there is pronounced variation in petal brightness, patch size, amount of dark markings, and contrast between patch and petal margin. We tested whether pollinators mediate selection on these color traits and on morphology (plant height, number of flowers, corolla size, spur length), and whether selection is consistent with facilitated or negative frequency‐dependent pollination. Pollinators mediated strong selection for increased petal brightness (Δβ_poll = 0.42) and contrast (Δβ_poll = 0.51). Pollinators also tended to mediate stabilizing selection on brightness (Δγ_poll = –0.27, n.s.) favoring the most common phenotype in the population. Selection for reduced petal brightness among hand‐pollinated plants indicated a fitness cost associated with brightness. The results demonstrate that flower color traits influence pollination success and seed production in A. morio, indicating that they affect attractiveness to pollinators, efficiency of pollen transfer, or both. The documented selection is consistent with facilitated pollination and selection for color convergence toward cooccurring rewarding species.     

Pollinator‐mediated selection on floral morphology: evidence for transgressive evolution in a derived hybrid lineage

Hybridization between closely related lineages is a mechanism that might promote substantive changes in phenotypic traits of descendants, resulting in transgressive evolution. Interbreeding between divergent but morphologically similar lineages can produce exceptional phenotypes, but the potential for transgressive variation to facilitate long‐term trait changes in derived hybrid lineages has received little attention. We compare pollinator‐mediated selection on transgressive floral traits in both early‐generation and derived hybrid lineages of the Piriqueta cistoides ssp. caroliniana complex. The bowl‐shaped flowers of morphotypes in this complex have similar gross morphologies and attract a common suite of small insect pollinators. However, they are defined by significant differences in characters that generate pollinator interest and visitation, including floral area and petal separation. In common garden experiments, patterns of pollen deposition in early‐generation recombinant hybrids indicate that Piriqueta's pollinators favour flowers with greater area and reduced petal separation. Changes in floral morphology in derived hybrid lineages are consistent with predictions from selection gradients, but the magnitude of change is limited relative to the range of transgressive variation. These results suggest that hybridization provides variation for evolution of divergent floral traits. However, the potential for extreme transgressive variants to contribute to phenotypic shifts may be limited due to reduced heritability, evolutionary constraints or fitness trade‐offs.     

The relative strength of different floral visitors driving floral evolution within a <Emphasis Type="Italic">Primula secundiflora</Emphasis> population

Floral visitor assemblages within plant populations are usually composed of different visitors, and the relative abundance of these visitors also varies. Therefore, identifying the relative strength of these floral visitors driving floral evolution within the population is an important step in predicting the evolutionary trajectory of floral traits. Using supplemental hand pollination and nectar-robbing exclusion treatments, we experimentally identified the relative strengths of legitimate pollinators (that visit flowers through the corolla tube entrance) and nectar robbers (that visit flowers by biting a hole in the corolla tube or using an existing hole) driving floral evolution within the Primula secundiflora population. We also estimated legitimate pollinator- and nectar robber-mediated selection separately for pin and thrum flowers. Both legitimate pollinators and nectar robbers mediated selection on pollination efficiency traits in P. secundiflora population. Legitimate pollinators mediated selection for wider corolla tubes, whereas nectar robbers mediated selection for longer corolla tubes. In addition, nectar robber-mediated selection on corolla tube length marginally varied between the pin and thrum flowers. Nectar robber mediated selection for longer corolla tube length in the pin flowers not in the thrum flowers. These results indicate that legitimate pollinators and nectar robbers within a population can drive differential evolutionary trajectories of floral traits.     

Induced mutations affecting pollinator choice in Mimulus lewisii (Phrymaceae)

The flowering plants are one of the most phenotypically varied and wide-ranging groups of organisms on earth, and yet, we have limited understanding of the contribution of animal pollinators to the diversification of floral form. To explore the interaction between variation in floral form and pollinator behavior, we observed the foraging behavior of bumblebees (Bombus impatiens) when presented with both wild-type Mimulus lewisii plants and each of three chemically induced single-locus mutants with altered floral phenotypes, including loss of the three lower petals, loss of nectar guides, and a change in petal color patterning. We found that each of the mutants attracted successful pollinator visits at just 29–80% of the rate relative to wild-type flowers, suggesting that effective recruitment of bumblebee pollinators requires the landing platform provided by the lower petals, and visual cues provided by the nectar guides and petal color pattern. Since single-locus recessive mutations are capable of ablating the lower petals, nectar guides, and color pattern, such changes in floral form provide insight into the driving forces behind plant adaptation.     

Pollinators and nectar robbers cause directional selection for large spur circle in Impatiens oxyanthera (Balsaminaceae)

Nectar spurs have an important role in floral evolution and plant–pollinator coadaptation. The flowers of some species possess spurs curving into a circle. However, it is unclear whether spur circle diameter is under direct selection pressure from different sources, such as pollinators and nectar robbers. In this study, we quantified selection on some floral traits, such as spur circle diameter in Impatiens oxyanthera (Balsaminaceae) using phenotypic selection analysis and compared the relative importance of pollinators and nectar robbers as selective agents using mediation analysis. The study showed that pollinators caused significant selection on corolla length, spur curvature and spur circle diameter while nectar robbers only imposed strong selection on spur circle diameter. Pollinators favored flowers with large corolla, curly spurs and large spur circle while nectar robbers preferred flowers with small spur circle. More pollinator visits resulted in higher female reproductive success, while robbery reduced female fitness. Conflicting selection on spur traits from pollinators and nectar robbers was not found. Mediation analysis showed that selection on floral traits through nectar robbing was stronger than selection through pollination. The results suggested that pollinators and nectar robbers jointly mediated the directional selection for large spur circle, and nectar robbers caused stronger selection than pollinators on floral traits.     

A test of phenotypic selection on petal form in the wild carnation,Dianthus inoxianus

Floral phenotypes are considered a product of pollinator‐mediated selection, which also has the side effect of decreasing floral variation within species. Correlates of flower visibility and function were studied in a carnation species (Dianthus inoxianus), which has crepuscular anthesis and scent‐based pollination by the hawkmoth Hyles livornica. We also assessed constancy of flower form in nature and in cultivation and, using fruit set as an estimate of plant relative fitness, tested whether the main pollinator exerted phenotypic selection on floral traits. Petal claw, which is roughly equivalent to the average depth at which an insect's proboscis must be inserted to reach nectar, was remarkably constant among wild plants (coefficient of variation 8%). In contrast, the area of the visible part of the petal, and the intensity of a coloured dot pattern on the petal was very variable (respectively CV = 34% and 102%). Cultivation in a common environment revealed significant variation among genotypes as regards petal area, degree of laciniation and extension of the dot pattern, but not petal claw length, which remained steady. Petal area, shape and colour did not affect relative fitness during the year of study, but plants with intermediate petal claws (i.e. floral tubes) set significantly more fruit. Results are compatible with low response of the main pollinator to variation in visual traits (petal area, laciniation, colour) and high responsiveness to variation in other aspects (tube length). Inconsistent phenotypic selection by pollinators may add to other causes of floral variation in the genus Dianthus, the causes of which are discussed.     

Pollinators exert selection on floral traits in a pollen-limited,narrowly endemic spring ephemeral

Premise

Floral traits are frequently under pollinator-mediated selection, especially in taxa subject to strong pollen-limitation, such as those reliant on pollinators. However, antagonists can be agents of selection on floral traits as well. The causes of selection acting on spring ephemerals are understudied though these species can experience particularly strong pollen-limitation. I examined pollinator- and antagonist-mediated selection in a narrowly endemic spring ephemeral, Trillium discolor.

Methods

I measured pollen limitation in T. discolor across two years and evaluated its breeding system. I compared selection on floral traits (display height, petal size, petal color, flowering time) between open-pollinated, and pollen-supplemented plants to measure the strength and mode of pollinator-mediated selection. I assessed whether natural levels of antagonism impacted selection on floral traits.

Results

Trillium discolor was self-incompatible and experienced pollen limitation in both years of the study. Pollinators exerted negative disruptive selection on display height and petals size. In one year, pollinator-mediated selection favored lighter petals but in the second year pollinators favored darker petals. Antagonist damage did not alter selection on floral traits.

Conclusions

Results demonstrate that pollinators mediate the strength and mode of selection on floral traits in T. discolor. Interannual variation in the strength, mode, and direction of pollinator-mediated selection on floral traits could be important for maintaining of floral diversity in this system. Observed levels of antagonism were weak agents of selection on floral traits.     

Natural selection on floral traits of female Silene dioica by a sexually transmitted disease

Floral traits endowing high reproductive fitness can also affect the probability of plants contracting sexually transmitted diseases. We explore how variations in floral traits influence the fitness of Silene dioica females in their interactions with pollinators carrying pollen or spores of the sterilizing anther-smut fungus Microbotryum violaceum. We collected healthy and infected plants in a highly diseased population and grew them under conditions that 'cure' infected individuals, and used standard regression methods to detect natural selection on floral traits. Narrow-sense heritabilities, coefficients of additive genetic variation (CV(A)) and genetic correlations among traits were estimated from paternal half-sib groups. Pollinator preferences imposed strong direct and directional selection on traits affecting female attractiveness and pollen-/spore-capturing abilities. Levels of additive genetic variance were high in these traits, suggesting that rapid responses to selection are possible. By considering our results in the light of spatial and temporal heterogeneity resulting from the colonization dynamics typical for this species, we suggest that the conflicting selective effects of pollen/spore loads lead to the maintenance of genetic variation in these traits.     

Oilseed rape (Brassica napus) as a resource for farmland insect pollinators: quantifying floral traits in conventional varieties and breeding systems

Oilseed rape (OSR; Brassica napus L.) is a major crop in temperate regions and provides an important source of nutrition to many of the yield‐enhancing insect flower visitors that consume floral nectar. The manipulation of mechanisms that control various crop plant traits for the benefit of pollinators has been suggested in the bid to increase food security, but little is known about inherent floral trait expression in contemporary OSR varieties or the breeding systems used in OSR breeding programmes. We studied a range of floral traits in glasshouse‐grown, certified conventional varieties of winter OSR to test for variation among and within breeding systems. We measured 24‐h nectar secretion rate, amount, concentration and ratio of nectar sugars per flower, and sizes and number of flowers produced per plant from 24 varieties of OSR representing open‐pollinated (OP), genic male sterility (GMS) hybrid and cytoplasmic male sterility (CMS) hybrid breeding systems. Sugar concentration was consistent among and within the breeding systems; however, GMS hybrids produced more nectar and more sugar per flower than CMS hybrid or OP varieties. With the exception of ratio of fructose/glucose in OP varieties, we found that nectar traits were consistent within all the breeding systems. When scaled, GMS hybrids produced 1.73 times more nectar resource per plant than OP varieties. Nectar production and amount of nectar sugar in OSR plants were independent of number and size of flowers. Our data show that floral traits of glasshouse‐grown OSR differed among breeding systems, suggesting that manipulation and enhancement of nectar rewards for insect flower visitors, including pollinators, could be included in future OSR breeding programmes.     

The structure of phenotypic variation in gender and floral traits within and among populations of Spergularia marina (Caryophyllaceae)

We sampled four wild populations of the highly autogamous Spergularia marina (Caryophyllaceae) in California to detect and to measure the magnitude of within- and among-population sources of phenotypic variation in gender and floral traits. From flowers and fruits collected from field and greenhouse-raised plants, we measured ovule number, seed number, mean seed mass, pollen production (greenhouse families only), mean pollen grain volume (greenhouse families only), anther number, anther/ovule ratio, pollen/ovule ratio (estimated using different flowers for pollen than for ovules; greenhouse families only), petal number, and petal size. Using greenhouse-raised genotypes, variation among maternal families nested within populations was evaluated for each trait to determine whether populations differ in the degree of maternally transmitted phenotypic variation. For each population, we used 15 greenhouse-raised maternal families to estimate the broad-sense heritability and genetic coefficient of variation of each floral trait. The magnitude and statistical significance of broad-sense heritability estimates were trait- and population-specific. Each population was characterized by a different combination of floral traits that expressed significant maternally transmitted (presumably genetic) variation under greenhouse conditions. Flowers representing two populations expressed low levels of maternally transmitted variation (three or fewer of nine measured traits exhibited significant maternal family effects on phenotype), while flowers representing the other two populations exhibited significant maternal family effects on phenotype for five or more traits. Our ability to detect statistically significant differences among the four populations depended upon the conditions under which plants were grown (field vs. greenhouse) and on the floral trait observed. Field-collected flowers exhibited significant differences among population means for all traits except anther number. Flowers sampled from greenhouse-raised maternal families differed among populations for all traits except ovule number, seed number, and petal size. We detected negligible evidence that genetic correlations constrain selection on floral traits in Spergularia marina.     

ECOLOGICAL COSTS OF PLANT RESISTANCE TO HERBIVORES IN THE CURRENCY OF POLLINATION

In this paper, we examine how ecological costs of resistance might be manifested through plant relationships with pollinators. If defensive compounds are incorporated into floral structures or if they are sufficiently costly that fewer rewards are offered to pollinators, pollinators may discriminate against more defended plants. Here we consider whether directional selection for increased resistance to herbivores could be constrained by opposing selection through pollinator discrimination against more defended plants. We used artificial selection to create two populations of Brassica rapa plants that had high and low myrosinase concentrations and, consequently, high and low resistance to flea beetle herbivores. We measured changes in floral characters of plants in both damaged and undamaged states from these populations with different resistances to flea beetle attack. We also measured pollinator visitation to plants, including numbers of pollinators and measures of visit quality (numbers of flowers visited and time spent per flower). Damage from herbivores resulted in reduced petal size, as did selection for high resistance to herbivores later in the plant lifetime. In addition, floral display (number of open flowers) was also altered by an interaction between these two effects. Changes in floral traits translated into overall greater use of low-resistance, undamaged plants based on total amount of time pollinators spent foraging on plants. Total numbers of pollinators attracted to plants did not differ among treatments; however, pollinators spent significantly more time per flower on plants from the low-resistance population and tended to visit more flowers on these plants as well. Previous work by other investigators on the same pollinator taxa has shown that longer visit times are associated with greater male and female plant fitness. Because initial numbers of pollinators did not differ between selection regimes, palatability and/or amount of rewards offered by high- and low-resistance populations are likely to be responsible for these patterns. During periods of pollinator limitation, less defended plants may have a selective advantage and pollinator preferences may mediate directional selection imposed by herbivores. In addition, if pollinator preferences limit seed set in highly defended plants, then lower seed set previously attributed to allocation costs of defense may also reflect greater pollinator limitation in these plants relative to less defended plants.     

Variation in nectar quality across 34 grassland plant species

• Floral nectar is considered the most important floral reward for attracting pollinators. It contains large amounts of carbohydrates besides variable concentrations of amino acids and thus represents an important food source for many pollinators. Its nutrient content and composition can, however, strongly vary within and between plant species. The factors driving this variation in nectar quality are still largely unclear.
• We investigated factors underlying interspecific variation in macronutrient composition of floral nectar in 34 different grassland plant species. Specifically, we tested for correlations between the phylogenetic relatedness and morphology of plants and the carbohydrate (C) and total amino acid (AA) composition and C:AA ratios of nectar.
• We found that compositions of carbohydrates and (essential) amino acids as well as C:AA ratios in nectar varied significantly within and between plant species. They showed no clear phylogenetic signal. Moreover, variation in carbohydrate composition was related to family-specific structural characteristics and combinations of morphological traits. Plants with nectar-exposing flowers, bowl- or parabolic-shaped flowers, as often found in the Apiaceae and Asteraceae, had nectar with higher proportions of hexoses, indicating a selective pressure to decelerate evaporation by increasing nectar osmolality.
• Our study suggests that variation in nectar nutrient composition is, among others, affected by family-specific combinations of morphological traits. However, even within species, variation in nectar quality is high. As nectar quality can strongly affect visitation patterns of pollinators and thus pollination success, this intra- and interspecific variation requires more studies to fully elucidate the underlying causes and the consequences for pollinator behaviour.

     

Pollen colour morphs take different paths to fitness

Colour phenotypes are often involved in communication and are thus under selection by species interactions. However, selection may also act on colour through correlated traits or alternative functions of biochemical pigments. Such forms of selection are instrumental in maintaining petal colour diversity in plants. Pollen colour also varies markedly, but the maintenance of this variation is little understood. In Campanula americana, pollen ranges from white to dark purple, with darker morphs garnering more pollinator visits and exhibiting elevated pollen performance under heat stress. Here, we generate an F2 population segregating for pollen colour and measure correlations with floral traits, pollen attributes and plant‐level traits related to fitness. We determine the pigment biochemistry of colour variants and evaluate maternal and paternal fitness of light and dark morphs by crossing within and between morphs. Pollen colour was largely uncorrelated with floral traits (petal colour, size, nectar traits) suggesting it can evolve independently. Darker pollen grains were larger and had higher anthocyanin content (cyanidin and peonidin) which may explain why they outperform light pollen under heat stress. Overall, pollen‐related fitness metrics were greater for dark pollen, and dark pollen sires generated seeds with higher germination potential. Conversely, light pollen plants produce 61% more flowers than dark, and 18% more seeds per fruit, suggesting a seed production advantage. Results indicate that light and dark morphs may achieve fitness through different means—dark morphs appear to have a pollen advantage whereas light morphs have an ovule advantage—helping to explain the maintenance of pollen colour variation.     

Floral antagonists counteract pollinator‐mediated selection on attractiveness traits in the hummingbird‐pollinated Collaea cipoensis (Fabaceae)

Pollinator‐mediated selection toward larger and abundant flowers is common in naturally pollen‐limited populations. However, floral antagonists may counteract this effect, maintaining smaller‐ and few‐flowered individuals within populations. We quantified pollinator and antagonist visit rates and determined a multiplicative female fitness component from attacked and non‐attacked flowers of the Brazilian hummingbird‐pollinated shrub Collaea cipoensis to determine the selective effects of pollinators and floral antagonists on flower size and number. We predicted that floral antagonists reduce the female fitness component and thus exert negative selective pressures on flower size and number, counteracting the positive effects of pollinators. Pollinators, mainly hummingbirds, comprised 4% of total floral visitation, whereas antagonist ants and bees accounted for 90% of visitation. Nectar‐robbers involved about 99% of floral antagonist visit rates, whereas florivores comprised the remaining 1%. Larger and abundant flowers increased both pollinator and antagonist visit rates and the female fitness component significantly decreased in flowers attacked by nectar‐robbers and florivores in comparison to non‐attacked flowers. We detected that pollinators favored larger‐ and many‐flowered individuals, whereas floral antagonists exerted negative selection on flower size and number. This study confirms that floral antagonists reduce female plant fitness and this pattern directly exerts negative selective pressures on flower size and number, counteracting pollinator‐mediated selection on floral attractiveness traits.     

Host adaptation in the anther smut fungus Ustilago violacea (Microbotryum violaceum): infection success,spore production and alteration of floral traits on two host species and their F1-hybrid

It is often assumed that host specialization is promoted by trade-offs in the performance of parasites on different host species, but experimental evidence for such trade-offs is scant. We studied differences in performance among strains of the anther smut fungus Ustilago violacea from two closely related host plant species, Silene alba and S. dioica, on progeny of (1) the host species from which they originated, (2) the alternative host species, and (3) inter-specific hybrids. Significant intra-specific variation in the pathogen was found for both infection success on a range of host genotypes (virulence) and components of spore production per infected host (aggressiveness) (sensu Burdon 1987). Strains did not have overall higher virulence on conspecifics of their host of origin than on strains from the heterospecific host, but they did have a significantly (c. 3 times) higher spore production per infected male host. This finding suggests that host adaptation may have evolved with respect to aggressiveness rather than virulence. The higher aggressiveness of strains on conspecifics of their host of origin resulted both from higher spore production per infected flower (spores are produced in the anthers), and greater ability to stimulate flower production on infected hosts. The latter indicates the presence of adaptive intraspecific variation in the ability of host manipulation. As transmission of the fungus is mediated by insects that are both pollinators of the host and vectors of the disease, we also assessed the effect of strains on host floral traits. Infection resulted in a reduction of inflorescence height, flower size, and nectar production per flower. Strains did not differ in their effect on nectar production, but infection with strains from S. alba resulted in a stronger reduction of inflorescence height and petal size on both host species. Vectors may therefore in principle discriminate among hosts infected by different strains and affect their efficiency of transmission. Contrary to assumptions of recent hypotheses about the role of host hybrids in the evolution of parasites, hybrids were not generally more susceptible than parental hosts. It is therefore unlikely that the rate of evolution of the pathogen on the parental species is slowed down by selection for specialization on the hybrids.     

Stabilizing selection on nectar concentration in wild Petunia axillaris,as revealed by genetic analysis of pollen dispersal

Most animal-pollinated plants produce nectar as a pollinator reward. Despite the main role that nectar plays in plant-pollinator interactions, the impact of natural variation in nectar traits on realized male fitness is poorly known. Here, we assessed this relation for a wild Petunia axillaris population using paternity-based direct selection gradient analysis, which allowed us also to infer pollen dispersal patterns. Because male fecundity may depend on other traits which could be associated with nectar characteristics (i.e. volume and concentration), we also considered selection on other key reproductive traits. The analysis revealed that P. axillaris was a strict outcrosser, but that successful pollination occurred mainly among neighbours. Individual plants varied greatly in their male fecundity. Nectar concentration, a key feature of nectar that determines its profitability, was subjected to stabilizing selection. Selection through male function also affected corolla area (positive directional selection), corolla tube length (negative directional selection), and floral display size (stabilizing selection), but none of these traits were phenotypically correlated with nectar characteristics. Because nectar concentration affects the ability and foraging efficiency of different flower visitors to feed on nectar, stabilizing selection may reflect either the preference of the most effective pollinators, or antagonistic selection driven by pollinators and non-pollinating nectar consumers.     

Foliar damage modifies floral attractiveness to pollinators in Alstroemeria exerens

Pollination is a requisite for sexual reproduction in plants and its success may determine the reproductive output of individuals. Pollinator preference for some floral designs or displays that are lacking or poorly developed in focal plants may constrain the pollination process. Foliar herbivory may affect the expression of floral traits, thus reducing pollinator attraction. Natural populations of the Andean species Alstroemeria exerens (Alstromeriaceae) in central Chile show high levels of foliar herbivory, and floral traits show phenotypic variation. In the present field study, we addressed the attractive role of floral traits in A. exerens and the effect of foliar damage on them. Particularly, we posed the following questions: (1) Is there an association between floral display and design traits and the number and duration of pollinator visits? and (2) Does foliar damage affect the floral traits associated with pollinator visitation? To assess the attractiveness of floral traits for pollinators, we recorded the number and duration of visits in 101 focal plants. To evaluate the effects of foliar damage on floral traits, 100 plants of similar size were randomly assigned to control or damage groups during early bud development. Damaged plants were clipped using scissors (50% of leaf area) and control plants were manually excluded from natural herbivores (<5% of leaf area eaten). During the peak of flowering, we recorded the number of open flowers, and estimated corolla and nectar guide areas. The number and duration of pollinator visits was statistically associated with floral design and display traits. Plants with larger displays, corollas and nectar guide areas received more visits. Visits lasted longer as display increases. In addition, foliar damage affected attractive traits. Damaged plants had fewer open flowers and smaller nectar guide areas. We conclude that foliar damage affects plant attractiveness for pollinators and hence may indirectly affect plant fitness.