Reconstructing the pollinator community and predicting seed set from hydrocarbon footprints on flowers

The measurement of insect visits to flowers is essential for basic and applied pollination ecology, but is often fraught with difficulty. Floral visitation is highly variable and observational studies are limited in scope due to the considerable time needed to acquire reliable data. Our study investigates whether the analysis of hydrocarbon residues (footprints) deposited by insects during flower visits allows the reconstruction of the visitor community and the prediction of seed set for large numbers of plants. In three consecutive years we recorded bumblebee visitation to wild plants of comfrey, Symphytum officinale, and later used gas chromatography/mass spectrometry (GC/MS) to quantify bumblebee-derived unsaturated hydrocarbons (UHCs) extracted from flowers. The UHCs washed from corollae were most similar to the tarsal UHC profile of the most abundant bumblebee species, Bombus pascuorum, in all 3 years. The species compositions of the bumblebee communities estimated from UHCs on flowers were also similar to those actually observed. There was a significant positive correlation between the observed number of visits by each of three bumblebee species (contributing 3–68% of the flower visits) and the estimated number of visits based on UHC profiles. Furthermore, significant correlations were obtained separately for workers and drones of two species. Seed set of comfrey plants was positively correlated with overall bumblebee visitation and the total amount of UHCs on flowers, suggesting the potential for pollen limitation. We suggest that quantifying cumulative footprint hydrocarbons provides a novel way to assess floral visitation by insects, and that this method can be used to predict seed set in pollen-limited plants.     

Floral display size in comfrey, Symphytum officinale L. (Boraginaceae): relationships with visitation by three bumblebee species and subsequent seed set

The fecundity of insect-pollinated plants may not be linearly related to the number of flowers produced, since floral display will influence pollinator foraging patterns. We may expect more visits to plants with more flowers, but do these large plants receive more or fewer visits per flower than small plants? Do all pollinator species respond in the same way? We would also expect foragers to move less between plants when the number of flowers per plant are large, which may reduce cross-pollination compared to plants with few flowers. We examine the relationships between numbers of inflorescence per plant, bumblebee foraging behaviour and seed set in comfrey, Symphytum officinale, a self-incompatible perennial herb. Bumblebee species differed in their response to the size of floral display. More individuals of Bombus pratorum and the nectar-robbing B.?terrestris were attracted to plants with larger floral displays, but B. pascuorum exhibited no increase in recruitment according to display size. Once attracted, all bee species visited more inflorescences per plant on plants with more inflorescences. Overall the visitation rate per inflorescence and seed set per flower was independent of the number of inflorescences per plant. Variation in seed set was not explained by the numbers of bumblebees attracted or by the number of inflorescences they visited for any bee species. However, the mean seed set per flower (1.18) was far below the maximum possible (4 per flower). We suggest that in this system seed set is not limited by pollination but by other factors, possibly nutritional resources.     

Spatio-temporal variation in the pollination mode of Buxus balearica (Buxaceae), an ambophilous and selfing species: mainland-island comparison

Mixed-pollination systems may allow plants to achieve stable seed production when unpredictable conditions cause variation in the relative success of different pollination modes. We studied variation in time (two years) and space (in five populations, three from an island and two from mainland) in the pollination mode of Buxus balearica , an ambophilous (i.e. pollinated by wind and insects) and selfing species distributed in the Mediterranean Basin, by means of direct observations and experimental manipulations (bagging with different material). The relative importance of each pollination mode differed among populations; however, levels of selfing and wind pollination were similar between island and mainland. Flowers of B. balearica were visited only by generalist insects, and species composition and abundance of flower visitors varied both in space and time. Frequency of insect visits to plants were not higher in mainland than island populations, although insects on the mainland were more diverse, visited a proportionally greater number of flowers, and remained longer on the plants than insects on the island. Frequency of insect visits was negatively correlated with flowering synchrony (all populations pooled) and was found to increase seed set in one of the mainland populations (that with highest frequency of insect visits and highest flower visitation rate). Fruit and seed mass were found to be not affected by pollination mode. Scarcity of pollinators in the island seems to have an effect on the pollination mode, although the greatest variation in breeding system was found at a more local scale.     

Size-dependent pollination efficiency in Anchusa officinalis (Boraginaceae): causes and consequences

Summary Bumblebees foraging on the self-incompatible Anchusa officinalis fly between near neighbour plants and between near neighbour inflorescences within plants. Although many-flowered plants attracted most bumblebees these plants received fewer visits on a per flower basis than smaller plants, and each bumblebee visited a smaller proportion of the flowers. The calculated effective visitation rate per flower was highest on plants of an intermediate size. If pollen-carryover was assumed to be limited the most efficient plant was predicted to be smaller since the proportion of fertilized flowers per bumblebee visit is expected to decrease further on the largest plants in relation to the total flower number. These predictions were tested by measuring fruit-set in the field. The percentage fruit-set decreased with plant size at all sizes that were investigated. That the most efficient plant was small indicates that pollen-carryover was indeed limited. However, the low percentage fruit-set associated with large size did not present a serious problem since the total estimated seed production per plant still increased with size. Selection favoring smaller plants may be low or absent in Anchusa.     

Pollination of Euphorbia dendroides by lizards and insects: Spatio-temporal variation in patterns of flower visitation

The patterns of flower visitation by lizards (Podarcis lilfordi, Lacertidae) and insects (mainly flies, bees and wasps) on the shrub Euphorbia dendroides, were studied in the island of Cabrera (Balearic Islands) during the flowering seasons of 1995 and 1996. Lizards act as true pollinators of the plant, moving large quantities of pollen within and among shrubs. To our knowledge, this is the first time that pollination by lizards has been empirically demonstrated. Variation in the quantitative component of pollination (frequency of visits × flower visitation rate) by the two groups of pollinators (lizards and insects) is documented at both spatial (within a plant population) and temporal scales (throughout the flowering season and between seasons). Variation in lizard density on a small spatial scale (within c. 200 m), presumably due to differences in vegetation cover, strongly affected their frequency of flower visitation. Insects were rather scarce, mainly because the plant flowers at a time (mid-March) when temperatures are still low. At the site where lizards were abundant, their frequency of flower visits was more than 3 times that of insects, they stayed on the shrubs about 3 times longer and visited about 8 times more cyathia per minute than did insects. Fruit and seed set were greater at this site, and this is attributed to the different frequency of flower visits by lizards, as shrubs are similar in size and produce similar amounts of cyathia in the two sites compared. Both, lizards and insects went more frequently to plants with large flower crops. However, flower crop was not associated with seed viability. We found no evidence for pollinator-mediated selection on plant traits related to fitness. Received: 28 August 1996 / Accepted: 26 February 1997     

Modification of bumblebee behavior by floral color change and implications for pollen transfer in <Emphasis Type="Italic">Weigela middendorffiana</Emphasis>

Flowers of Weigela middendorffiana change the color from yellow to red. The previous study revealed that red-phase flowers no longer have sexual function and nectar, and bumblebees selectively visit yellow-phase flowers. The present study examined how retaining color-changed flowers can regulate the foraging behavior of bumblebees and pollen transport among flowers within (geitonogamous pollination) and between (outcrossing pollination) plants and how the behavior is influenced by display size (i.e., number of functional flowers) and visitation frequency. The visitation frequencies of bumblebees to plants and successive flower probes within plants were observed in the field using plants whose flower number and composition of the two color-phase flowers had been manipulated. To evaluate pollination efficiency over multiple pollinator visits, a pollen transport model was constructed based on the observed bumblebee behavior. In the simulation, three flowering patterns associated with display size and existence of color-changed flowers were postulated as follows: Type 1, large display (100 functional flowers) and no retention of color-changed flowers; Type 2, small display (50 functional flowers) and retention of color-changed flowers (50 old flowers), and; Type 3, large display (100 functional flowers) and retention of color-changed flowers (100 old flowers). Color-changed flowers did not contribute to increasing bumblebee attraction at a distance but reduced the number of successive flower probes within plants. Comparisons of pollen transfer between Types 1 and 3 revealed that the retention of color-changed flowers did not influence the total amount of pollen exported when pollinator visits were abundant (>100 visits) but decreased geitonogamous pollination. Comparisons between Types 2 and 3 revealed that the discouragement effect of floral color change on successive probes accelerated in plants with a large display size. Overall, the floral color change strategy contributed to reduce geitonogamous pollination, but its effectiveness was highly sensitive to display size and pollinator frequency.     

Contribution of small insects to pollination of common buckwheat, a distylous crop

Crop pollination by animals is an essential ecosystem service. Among animal-pollinated crops, distylous plants strongly depend on animal pollination. In distylous pollination systems, pollinator species are usually limited, although flowers of some distylous plants are visited by diverse animals. We studied the pollination biology of common buckwheat ( Fagopyrum esculentum ), a distylous crop mainly pollinated by honeybees and visited by many insect species, to evaluate the effects of non-honeybee species on pollination services. We focused on insects smaller than honeybees to determine their contribution to pollination. We applied pollination treatments with bags of coarse mesh to exclude flower visits by honeybees and larger insects and compared the seed set of bagged plants with that of untreated plants for pin and thrum flower morphs. We found a great reduction of seed set only in bagged pin flowers. We also confirmed that small insects, including ants, bees, wasps and flies, carried pin-morph pollen. These small insects transfer pollen from the short anthers of pin flowers to the short styles of thrum flowers, leading to sufficient seed set in thrum flowers. Consequently, small, non-honeybee insects have the potential to maintain at least half of the yield of this honeybee-dependent distylous crop.     

Sources of variation in pollinator contribution within a guild: the effects of plant and pollinator factors

Among plants visited by many pollinator species, the relative contribution of each pollinator to plant reproduction is determined by variation in both pollinator and plant traits. Here we evaluate how pollinator movement among plants, apparent pollen carryover, ovule number, resource limitation of seed set, and pollen output affect variation in contribution of individual pollinator species to seed set in Lithophragma parviflorum (Saxifragaceae), a species visited by a broad spectrum of visitors, including beeflies, bees and a moth species. A previous study demonstrated differences among visitor species in their single-visit pollination efficacy but did not evaluate how differences in visitation patterns and pollen carryover affect pollinator efficacy. Incorporation of differential visitation patterns and pollen carryover effects —commonly cited as potentially important in evaluating pollinator guilds — had minor effects (0–0.6% change) on the estimates of relative contribution based on visit frequency and single-visit efficacy alone. Beeflies visited significantly more flowers per inflorescence than the bees and the moth. Seed set remained virtually constant during the first three visited flowers for beeflies and larger bees, indicating that apparent pollen carryover did not reduce per-visit efficacy of these taxa. In contrast, Greya moth visits showed a decrease in seed set by 55.4% and the smaller bees by 45.4% from first to second flower. The larger carryover effects in smaller bees and Greya were diminished in importance by their small overall contribution to seed set. Three variable plant traits may affect seed set: ovule number, resource limitation on seed maturation, and pollen output. Ovule number per flower declined strongly with later position within inflorescences. Numbers were much higher in first-year greenhouse-grown plants than in field populations, and differences increased during 3 years of study. Mean pollen count by position varied 7-fold among flowers; it paralleled ovule number variation, resulting in a relatively stable pollen:ovule ratio. Resource limitation of seed set increased strongly with later flowering, with seed set in hand-pollinated flowers ranging from 66% in early flowers to 0% in the last two flowers of all plants. Variation in ovule number and resource limitation of seed maturation jointly had a strong effect on the number of seeds per flower. Visitation to early flowers had the potential to cause more seed set than visitation to later flowers. Overall, the most important sources of variation to seed production contribution were differences among pollinators in abundance and absolute efficacy (ovules fertilized on a single visit) and potentially differential phenology among visitor species. These effects are likely to vary among populations and years.     

THE EVOLUTION OF INSECT-MEDIATED SELF-POLLINATION FROM AN OUTCROSSING SYSTEM IN CALYPTRIDIUM (PORTULACACEAE)

Calyptridium monospermum (Portulacaceae) comprises two biological races visited by different groups of insects. One race, pollinated by a bumblebee, Bombus vosnesenskii (Apidae), is outcrossing, while the other race is pollinated casually by a variety of other insects whose visits result in insect-mediated self-pollination and geitonogamy within the inflorescence. Insect-mediated selfing is also the mode of pollination in the closely related species, C. umbellatum. While all plants of both species are self-compatible, insect visitation is necessary for good seed set. Selfing populations of C. monospermum exhibit many floral adaptations for insect-mediated self-pollination which parallel those of the selfing C. umbellatum, while retaining vegetative characteristics typical of C. monospermum. These floral traits involve flower color and odor, style and stamen orientation, size of stigmatic surfaces, number of pollen grains produced, density of open flowers on inflorescences, and presence or absence of protogyny. Artifical pollination experiments show that these floral adaptations are effective in increasing the probability of insect-mediated self-pollination. Insect-mediated selfing in C. umbellatum and in some populations of C. monospermum is encouraged and probably made necessary by the inconstant pollinating behavior of the insects which visit these plants, as shown by the large percentage of non-Calyptridium pollen recovered from the bodies of insects captured while visiting these plants.     

The effects of floral display on pollinator visitation vary among populations ofPhacelia linearis (Hydrophyllaceae)

Summary Individual plants in gynodioecious populations ofPhacelia linearis (Hydrophyllaceae) vary in flower gender, flower size, and flower number. This paper reports the effects of variation in floral display on the visitation behaviour of this species' pollinators (mainly pollen-collecting solitary bees) in several natural and three experimental plant populations, and discusses the results in terms of the consequences for plant fitness. The working hypotheses were: (1) that because female plants do not produce pollen, pollen-collecting insects would visit hermaphrodite plants at a higher rate than female plants and would visit more flowers per hermaphrodite than per female; and (2) that pollinator arrival rate would increase with flower size and flower number, the two main components of visual display. These hypotheses were generally supported, but the effects of floral display on pollinator visitation varied substantially among plant populations. Hermaphrodites received significantly higher rates of pollinator arrivals and significantly higher rates of visits to flowers than did females in all experimental populations. Flower size affected arrival rate and flower visit rate positively in natural populations and in two of the three experimental populations. The flower size effect was significant only among female plants in one experimental population, and only among hermaphrodites in another. The effect of flower number on arrival rate was positive and highly significant in natural populations and in all experimental populations. In two out of three experimental populations, insects visited significantly more flowers per hermaphrodite than per female and visited more flowers on many-flowered plants than on few-flowered plants, but neither effect was detected in the third experimental population. Because seed production is not pollen-limited in this species, variation in pollinator visitation behaviour should mainly affect the male reproductive success of hermaphrodite plants. These findings suggest that pollinator-mediated natural selection for floral display inP. linearis varies in space and time.     

Pollination biology of Prunus mahaleb L.: deferred consequences of gender variation for fecundity and seed size

This study describes the reproductive biology of Prunus mahaleb, a rosaceous treelet, in a southeastern Spanish population. The species is gynodioecious with 55.4% of the plants being male-fertile and 44.6% presenting non-functional, shrunken anthers with no pollen, and behaving as functional females. Individual trees produced the same flower morph in four consecutive study years. Fruit set in bagged inflorescences was absent in male-sterile trees and was very reduced in male-fertiles. Open-pollinated flowers of male-fertiles showed greater fruit set (x?= 29.1%) than male-steriles (x?= 25.3%). Average fruit set in three experimental treatments (control, selfed, and crossed) were 29.05%, 41.9% and 38.6%, respectively, for male-fertile trees; those for male-steriles were 25.3%, 0% and 39.2%, respectively. Seeds from male-sterile plants were heavier (71.33 mg) than those of male-fertile plants (66.05 mg) but did not differ in germination ability. Male-fertile and male-sterile trees differ significantly in average nectar concentration (73.5% and 55.9%, respectively) and nectar volume secreted/flower and day (0.142 μl and 0.171 μl, respectively). A total of 41 species of insect flower visitors were recorded. Calliphorid and tachinid flies (41.97% of total visits), and andrenid bees (30.30%), were the most frequent visitors. The diversity of insect visitors was greater in male-fertile trees. Bees and flies accounted for 50.5% and 49.5%, respectively of total visits to male-fertile trees. Flies were far more frequent than bees at male-sterile flowers (76.9% and 23.1%, respectively). Male-sterile trees received higher average visitation (47 insects/census) than male-fertile trees (32 insects/census). Individual trees. showed a relative constancy of the fecundity rankings between years. Fruit production was significantly higher in male-sterile trees, with a four-year average of 6558 fruits, in contrast with male-fertile trees which yielded 4670 fruits. This 1.77-fold difference in favour of male-sterile trees over malefertiles was not compensated by the greater fruit set of the latter. This is attributable to the lower outcrossing rate of male-fertiles, estimated as 52.68% from seed mass data. In addition, greater seed mass of female progeny, and higher visitation rate by insects might explain the maintenance of this polymorphism. These results have far-reaching implications for the demography and seed-dispersal ecology of this endozoochorus species.     

Do flowers wave to attract pollinators? A case study with Silene maritima

To answer the question whether flowers wave to attract pollinators, we determine: (1) the heritability of floral mobility; (2) whether wavy flowers attract more insects; (3) does the duration of pollination affect seed set; and (4) the relationship between seed set and floral mobility. The pollination ecology of Silene maritima was investigated. Flowers on stalks of different waviness were used to investigate the effect of floral movement on pollinator visits. There is heritable variation in both direct and indirect estimates of floral mobility. The highest insect total visitation times were associated with medium length thin stalks that were visited more frequently and by more insect species. Although mean individual visit durations were less than those of less mobile flowers, this was compensated for by increased visits. Observations of controlled pollinations show that when the visit times are low, so is seed set and therefore low and high mobility flowers might suffer from reduced fitness. Combining these observations provides a mechanism that could be driving stabilizing selection for flower stalk traits, with a trade-off applying between waving to attract pollinators and not being too mobile as to prevent effective pollination. Further evidence for stabilizing selection is provided by the relationship observed in the field between seed set and floral mobility where the highest levels of fitness was associated with intermediate levels of floral waving.     

The presence of co-flowering species facilitates reproductive success of Pedicularis monbeigiana (Orobanchaceae) through variation in bumble-bee foraging behaviour

Background and Aims The presence of co-flowering species can alter pollinator foraging behaviour and, in turn, positively or negatively affect the reproductive success of the focal species. Such interactions were investigated between a focal species, Pedicularis monbeigiana, and a co-flowering species, Vicia dichroantha, which was mediated by behaviour alteration of the shared bumble-bee pollinator. Methods Floral display size and floral colour change of P. monbeigiana were compared between pure (P. monbeigiana only) and mixed (P. monbeigiana and V. dichroantha) plots in two populations. Pollinator visitation rates, interspecific floral switching and successive within-plant pollinator visits were recorded. In addition, supplemental pollination at plant level was performed, and the fruit set and seed set were analysed in pure and mixed plots with different densities of P. monbeigiana. Key Results Pollinator visitation rates were dramatically higher in mixed plots than in pure plots. The higher pollinator visitation rates were recorded in both low- and high-density plots. In particular, successive flower visits within an individual plant were significantly lower in mixed plots. Supplemental pollination significantly increased fruit set and seed set of individuals in pure plots, while it only marginally increased seed set per fruit of plants in mixed plots. Conclusions The presence of V. dichroantha can facilitate pollination and increase female reproductive success of P. monbeigiana via both quantity (mitigating pollinator limitation) and quality (reducing geitonogamy) effects. This study suggests that successive pollinator movements among flowers within a plant, as well as pollinator visitation rates and interspecific flower switching, may be important determinants of the direction and mechanisms of interaction between species.     

Size and sex of floral displays affect insect visitation rates in the dioecious Australian alpine herb, Aciphylla glacialis

Differences in floral display size between male and female plants in dioecious species are often considered to be the result of competition within and between male and female plants to attract insects, although this hypothesis is rarely tested. In a field study of the alpine dioecious herb Aciphylla glacialis , insect visitation rates were found to increase with increasing floral display, as predicted. Dipterans were the most common visitors to male and female inflorescences, both in terms of individuals visits (2083 out of 2581 total visits by insects), and species number (20 morphospecies, potentially 22 species). Larger male inflorescences attracted 1.7 times as many insects in total, and 3.1 times more visits by the most common flower visitor, Musca vetustissima than did female inflorescences, but equal number of visits by the next most common flying visitors, Poecilohetaerus aquilius and Tephritis poenia. There was no difference in the amount of time M. vetustissima individuals spent on male and female inflorescences, even though male inflorescences have more flowers. Larger displays (four and eight inflorescences) of either sex attracted more visits than smaller ones (one inflorescence), although the rate of increase in visitation rate with increasing floral display declined. The response of insects to increasing floral display does not support the predicted escalating or proportionate increasing rate of visitation of some models for the evolution of dimorphism in floral displays in dioecious plants.     

Floral display, pollinator visitation and reproductive success in the dioecious perennial herb Wurmbea dioica (Liliaceae)

Floral traits that increase attractiveness to pollinators are predicted to evolve through selection on male function rather than on female function. To determine the importance of male-biased selection in dioecious Wurmbea dioica, we examined sexual dimorphism in flower size and number and the effects of these traits on pollinator visitation and reproductive success of male and female plants. Males produced more and larger flowers than did females. Bees and butterflies responded to this dimorphism and visited males more frequently than females, although flies did not differentiate between the sexes. Within sexes, insect pollinators made more visits to and visited more flowers on plants with many flowers. However, visits per flower did not vary with flower number, indicating that visitation was proportional to the number of flowers per plant. When flower number was experimentally held constant, visitation increased with flower size under sunny but not overcast conditions. Flower size but not number affected pollen removal per flower in males and deposition in females. In males, pollen removal increased with flower size 3 days after flowers opened, but not after 6 days when 98% of pollen was removed. Males with larger flowers therefore, may have higher fitness not because pollen removal is more complete, but because pollen is removed more rapidly providing opportunities to pre-empt ovules. In females, pollen deposition increased with flower size 3 days but not 6 days after flowers opened. At both times, deposition exceeded ovule production by four-fold or more, and for 2 years seed production was not limited by pollen. Flower size had no effect on seed production per plant and was negatively related to percent seed set, implying a tradeoff between allocation to attraction and reproductive success. This indicates that larger flower size in females is unlikely to increase fitness. In both sexes, gamete production was positively correlated with flower size. In males, greater pollen production would increase the advantage of large flowers, but in females more ovules may represent a resource cost. Selection to increase flower size and number in W. dioica has probably occurred through male rather than female function. Received: 15 June 1997 / Accepted: 12 February 1998     

Effects of patch size and density on flower visitation and seed set of wild plants: a pan-European approach

1.  Habitat fragmentation can affect pollinator and plant population structure in terms of species composition, abundance, area covered and density of flowering plants. This, in turn, may affect pollinator visitation frequency, pollen deposition, seed set and plant fitness.
2.  A reduction in the quantity of flower visits can be coupled with a reduction in the quality of pollination service and hence the plants' overall reproductive success and long-term survival. Understanding the relationship between plant population size and/or isolation and pollination limitation is of fundamental importance for plant conservation.
3.  We examined flower visitation and seed set of 10 different plant species from five European countries to investigate the general effects of plant populations size and density, both within (patch level) and between populations (population level), on seed set and pollination limitation.
4.  We found evidence that the effects of area and density of flowering plant assemblages were generally more pronounced at the patch level than at the population level. We also found that patch and population level together influenced flower visitation and seed set, and the latter increased with increasing patch area and density, but this effect was only apparent in small populations.
5.   Synthesis. By using an extensive pan-European data set on flower visitation and seed set we have identified a general pattern in the interplay between the attractiveness of flowering plant patches for pollinators and density dependence of flower visitation, and also a strong plant species-specific response to habitat fragmentation effects. This can guide efforts to conserve plant–pollinator interactions, ecosystem functioning and plant fitness in fragmented habitats.     

Pollination ecology of Silene acutifolia (Caryophyllaceae): floral traits variation and pollinator attraction

BACKGROUND AND AIMS: The floral display influences the composition of pollinators interacting with a plant species. Geographic and temporal variation in pollinator composition complicates the understanding of the evolutionary consequences of floral display variation. This paper analyses the relationships between Silene acutifolia, a hermaphroditic perennial herb, and its pollinators, based on field studies in the north-west of Spain. METHODS: Studies were conducted over three years (1997-1999). Firstly, the main pollinators of this species were determined for two years in one population. Secondly, pollen limitation in fruit and seed production was analysed by supplementary hand pollinations, and counting the pollen grains and tubes growing in styles for two different-sized populations. Finally, the effect of flower size and number on the rate of visitation and total seed number was examined for 15 marked plants. RESULTS AND CONCLUSIONS: The primary pollinators were long-tongued insects, including Hymenoptera, Lepidoptera and Diptera, but the composition and visitation frequencies differed between years. Pollen limitation occurred in one of the years of study. There was between-population variation in the number of pollen grains and pollen tubes found in styles, suggesting pollen limitation in one population. Overall, pollinators visited plants with more open flowers more frequently, and pollinated more flowers within these plants. Conversely, petal and calyx sizes had no effect on insect visitation. Plants with higher rates of visits produced higher number of seeds, suggesting that pollinator-mediated limitation of seed and fruit production may be important in some years.     

Pollinators shift to nectar robbers when florivory occurs,with effects on reproductive success in Iris bulleyana (Iridaceae)

• Studies have indicated that florivory and nectar robbing may reduce reproductive success of host plants. However, whether and how these effects might interact when plants are simultaneously attacked by both florivores and nectar robbers still needs further investigation.
• We used Iris bulleyana to detect the interactions among florivory, nectar robbing and pollination, and moreover, their effects on plant reproductive success. Field investigations and hand‐pollination treatments were conducted on two experimental plots from a natural population, in which Experimental plot was protected from florivores and Control plot was not manipulated.
• The flower calyx was bitten by sawflies to consume the nectary, and three bumblebee species were pollinators. In addition, the short‐tongued pollinator, Bombus friseanus, was the only robber when there was a hole made by a sawfly. The bumblebee had significantly shortened flower handling time when robbing, as compared to legitimate visits. Pollinator visitation and seed production decreased significantly in damaged flowers. However, seed production per flower after supplementary hand‐pollination did not differ significantly between damaged and undamaged flowers. Compared to the Experimental plot, bumblebees visited fewer flowers per plant in a foraging bout in the Control plot.
• The flowers damaged by florivory allowed B. friseanus to shift to a nectar robber. Florivory and nectar robbing collectively decreased plant reproductive success by consuming nectar resources, which may reduce attractiveness to pollinators of the damaged flowers. However, the changes in pollinator behaviour might be beneficial to the plant by reducing the risk of geitonogamous mating.

     

Arbuscular mycorrhizal fungi influence visitation rates of pollinating insects

Abstract.  1. Arbuscular mycorrhizal (AM) fungi can increase a number of plant traits to which pollinating insects are known to respond. These include total plant size, flower number, flower size, and amount of pollen produced.
2. It was hypothesised that these effects would lead to a different visitation rate of pollinating insects on mycorrhizal and non-mycorrhizal plants. To test this idea, three species of annual plants ( Centaurea cyanus , Tagetes erecta , and Tagetes patula ) were grown with and without AM fungi and the visits by pollinating insects were recorded over a 2-month period.
3. In all three species, mycorrhizal plants experienced a greater number of pollinator visits per flower per unit time. Diptera and Hymenoptera were the predominant insects and the latter order showed the strongest response.
4. Here, it is suggested that mycorrhizal fungi increase floral visitation rates by insects, but that the mechanism varies from one plant species to another. In C. cyanus , it appears to be due to flower number per plant, in T. patula it is individual inflorescence size, and in T. patula it is nectar standing crop per inflorescence.     

Role of honey bees (Hymenoptera: Apidae) in the pollination biology of a California native plant, Triteleia laxa (Asparagales: Themidaceae)

A central focus of pollination biology is to document the relative effectiveness of different flower visitors as pollinators. Ongoing research seeks to determine the role that introduced honey bees (Apis mellifera L.) play in the pollination of both invasive and native plants. Here we report on the importance of A. mellifera as pollinators of a California native plant, Triteleia laxa Bentham. In observation plots and transect censuses, A. mellifera overwhelmingly dominated the T. laxa flower visitor assemblage. We believe the proximity to agriculture, where A. mellifera density is higher relative to areas far from agriculture, contributes to the discrepancy between A. mellifera abundance at the two sites. Although A. mellifera were inferior flower visitors qualitatively (visited less flowers per minute), they were the most frequent interactors with flowers. Furthermore, the proportion of visits to flowers on the same plant among flower visitor species did not differ, suggesting a general mechanism by which insects forage at T. laxa flowers and that A. mellifera do not cause more deleterious geitonogamy than do native pollinators. Flower visitation rates as a function of floral display size did not differ between A. mellifera and other flower visitors. The difference in the magnitude of flower visitation (largely by A. mellifera) between sites is consistent with a difference in seed set between sites. These results suggest that non-native A. mellifera bees can play an important role in the pollination of native plant species.