Effects of nectar robbing on nectar dynamics and bumblebee foraging strategies in Linaria vulgaris (Scrophulariaceae)

Differences in morphology among bumblebee species sharing a nectar resource may lead to variation in foraging behaviour and efficiency. Less efficient bumblebees might opportunistically switch foraging strategies from legitimate visitation to secondary robbing when hole-biting primary robbers are present. We observed various aspects of pollination and nectar robbing ecology of Linaria vulgaris in the Colorado Rocky Mountains, with emphasis on the role of bumblebee proboscis length. Bees can extract nectar from a nectar spur legitimately, by entering the front of the flower, or illegitimately, by biting or reusing holes in the spur. Although L. vulgaris flowers are apparently adapted for pollination by long-tongued bees, short-tongued bees visited them legitimately for trace amounts of nectar but switched to secondary robbing in the presence of primary robbers. Longer-tongued bees removed more nectar in less time than did shorter-tongued bees, and were less likely to switch to secondary robbing even when ∼100% of flowers had been pierced. As the proportion of robbed flowers in the population increased, the relative number of legitimate visits decreased while the relative number of robbing visits increased. Robbing decreased nectar standing crop and increased the proportion of empty flowers per inflorescence. Despite these potentially detrimental effects of robbers, differences in inflorescence use among robbers and pollinators, and the placement of holes made by primary robbers, may mitigate negative effects of nectar robbing in L. vulgaris . We discuss some of the reasons that L. vulgaris pollination ecology and growth form might temper the potentially negative effect of nectar robbing.     

How long to stay on a plant: the response of bumblebees to encountered nectar levels

This field study shows that the number of flowers visited per bee per plant (Anchusa officinalis) increases with the instantaneous nectar level at the plant. Observations during the season showed that a bee visits more flowers per plant of given nectar level, the lower the overall mean nectar level in the study area. These results agree with predictions from a model based on the ‘marginal value theorem’, but with assumptions and constraints adapted for nectar-foraging bees. It suggests that bumblebees assess the nectar level at a plant by sampling one or a few flowers, which is possible because within-plant nectar volumes are correlated. The bees compare encountered gains to an optimal plant switching threshold equal to the overall mean nectar level and leave an unrewarding plant as soon as possible, but continue to visit the flowers on a rewarding plant. However, the bees leave before having visited all flowers due to a searching constraint. The bees’ response to plant nectar levels results in systematic flower visitation, because visitation to recently depleted flowers is reduced, which reduces the variation of the inter-visit time per flower. Systematic flower visitation implies that the overall mean encountered gain per flower is higher than the overall mean standing crop, as predicted by a model of systematic foraging. However, the sampling and searching constraints on the bees’ response to plant nectar levels increase the variation of the inter-visit time per flower, and thereby limit the degree of systematic flower visitation and the effect on the mean encountered gain.     

Flowers morphology and nectar concentration determine the preferred food source of stingless bee,Heterotrigona itama

Heterotrigona itama is a stingless bee species from Meliponini tribe. The bee collects nectar, pollen and resin to produce honey, bee bread, and propolis. The bee is also known to visit and collect nectar from various types of flowers but there are limited studies on why this species of bee prefers to visit certain types of flowers. This study was conducted to identify the nectar concentration in selected flowers favoured by H. itama and the relationship between the bee and the morphology of the flowers. Nectar was obtained from different species of flowers and the concentrations were measured using a digital refractometer. The tube length of each flower species and the tongue length of the bees were also measured. The results revealed that flowers preferred by H. itama have high nectar concentrations. The tube lengths of the preferred flowers were between 2.0 and 4.0 mm, which is compatible with the tongue length of the bee. This study revealed that both nectar concentration and flower morphology are important factors for the bees in choosing their food sources. The results from this study will benefit the beekeepers in the identification of flowers that should be planted in their farms to improve stingless bee beekeeping activities. Understanding the relationship between the bees and their flower preferences could also help us to understand the importance of conserving both the bee colonies and the various species of flowering plants to ensure the sustainability of flora and fauna in the ecosystem.     

Evidence for mutualism between a flower-piercing carpenter bee and ocotillo: use of pollen and nectar by nesting bees

Abstract.

• 1 Carpenter bees (Xylocopa californica arizonensis) in west Texas, U.S.A., gather pollen and ‘rob’ nectar from flowers of ocotillo (Fouquieria splendens). When common, carpenter bees are an effective pollen vector for ocotillo. We examined ocotillo's importance as a food source for carpenter bees.
• 2 The visitation rate of carpenter bees to ocotillo flowers in 1988 averaged 0.51 visits/flower/h and was 4 times greater than that of queen bumble bees (Bombus pennsylvanicus sonorus), the next most common visitor. Nectar was harvested thoroughly and pollen was removed from the majority of flowers. Hummingbird visits were rare.
• 3 Pollen grains from larval food provisions were identified from sixteen carpenter bee nests. On average, 53% of pollen grains sampled were ocotillo, 39% were mesquite (Prosopis glandulosa), and 8% were Zygophyllaceae (Larrea tridentata or Guaiacum angustifolium). Carpenter bee brood size averaged 5.8 per nest.
• 4 We measured the number of flowers, and production of pollen and nectar per flower by mature ocotillo plants, as well as the quantity of pollen and sugar in larval provisions. An average plant produced enough pollen and nectar sugar to support the growth of eight to thirteen bee larvae. Ocotillo thus has the potential to contribute significantly to population growth of one of its key pollinators.
• 5 Although this carpenter bee species, like others, is a nectar parasite of many plant species, it appears to be engaged in a strong mutualism with a plant that serves as both a pollen and as a nectar source during carpenter bee breeding periods.

     

Pollinator foraging modifies nectar sugar composition in Helleborus foetidus (Ranunculaceae):An experimental test

We experimentally tested the hypothesis that the extensive within-plant variation of nectar sugar composition in Helleborus foetidus (Ranunculaceae) and other species results from differences between flowers and nectaries in pollinator visitation history. Experiments were conducted to mimic single-nectary visits by wild-caught individuals of the main bee pollinators of H. foetidus, which were assayed for their capacity to modify the sugar composition of natural and artificial nectar. Experimental nectar probing with bee mouthparts induced extensive changes in proportional sugar composition 48 h after treatment, and bee taxa differed widely in their effects. Nectar probing by Andrena, medium-sized Anthophoridae, Apis mellifera, and Lasioglossum had no subsequent effects on nectar sugar composition, while probing by Bombus terrestris and B. pratorum induced an extensive reduction in percentage sucrose, a marked increase in percentage fructose, and a slight increase in percentage glucose. Results support the hypothesis that stochastic variations among flowers or nectaries in the taxonomic identity of recent visitors and their relative visitation frequencies may eventually generate very small-scale mosaics in nectar sugar composition. Changes in nectar sugar composition following bumblebee probing may be the consequence of nectar contamination with pollinator-borne nectarivorous yeasts.     

Visitation rate of pollinators and nectar robbers to the flowers and inflorescences of Tabebuia aurea (Bignoniaceae): effects of floral display size and habitat fragmentation

Large floral displays favour pollinator attraction and the import and export of pollen. However, large floral displays also have negative effects, such as increased geitonogamy, pollen discounting and nectar/pollen robber attraction. The size of the floral display can be measured at different scales (e.g. the flower, inflorescence or entire plant) and variations in one of these scales may affect the behaviour of flower visitors in different ways. Moreover, the fragmentation of natural forests may affect flower visitation rates and flower visitor behaviour. In the present study, video recordings of the inflorescences of a tree species (Tabebuia aurea) from the tropical savannah of central Brazil were used to examine the effect of floral display size at the inflorescence and tree scales on the visitation rate of pollinators and nectar robbers to the inflorescence, the number of flowers approached per visit, the number of visits per flower of potential pollinators and nectar robbers, and the interaction of these variables with the degree of landscape disturbance. Nectar production was quantified with respect to flower age. Although large bees are responsible for most of the pollination, a great diversity of flower insects visit the inflorescences of T. aurea. Other bee and hummingbird species are highly active nectar robbers. Increases in inflorescence size increase the visitation rate of pollinators to inflorescences, whereas increases in the number of inflorescences on the tree decrease visitation rates to inflorescences and flowers. This effect has been strongly correlated with urban environments in which trees with the largest floral displays are observed. Pollinating bees (and nectar robbers) visit few flowers per inflorescence and concentrate visits to a fraction of available flowers, generating an overdispersed distribution of the number of visits per inflorescence and per flower. This behaviour reflects preferential visits to young flowers (including flower buds) with a greater nectar supply.     

西方蜜蜂和兰州熊蜂在设施桃园的访花偏好性比较

【目的】为了比较西方蜜蜂 Apis mellifera 和兰州熊蜂 Bombus lantschouensis 在设施桃园内对不同时期桃花的访花偏好性、以及这种偏好性与花粉活力和采集花粉花蜜之间的关系。【方法】记录2种蜂在温室桃园内访问早期花、中期花和晚期花的比例,测定桃花不同时期的花粉活力以及2种蜂携带的花粉活力,观察2种蜂采集花蜜和采集花粉的成功率,统计2种蜂访花过程中桃花所处的枝条数及植株数。【结果】桃花不同时期的花粉活力差异显著,早期花花药未开裂,花粉未释放,中期花花粉活力（58.3%）显著高于晚期花花粉活力（34.2%）（P<0.01）;西方蜜蜂更加偏好访问中期花,对中期花的访问率高达75.3%,显著高于兰州熊蜂对中期花的访问率（49.2%）（P<0.01）;西方蜜蜂携带的花粉活力（92.1%）显著高于兰州熊蜂携带的花粉活力（72.9%）,但是西方蜜蜂采集花粉和采集花蜜的成功率均低于兰州熊蜂（P<0.01）;在访问一定数量的桃花过程中,兰州熊蜂在设施桃园内访问的枝条数和植株数较多,分布范围较广（P<0.01）。【结论】和兰州熊蜂相比,西方蜜蜂对活力花粉的辨别能力更强,更加偏好访问花粉活力较高的花朵,这种偏好性导致其采集花粉花蜜的成功率降低。     

Spatial distribution of nectar production in a natural Echium vulgare population: Implications for pollinator behaviour

The distribution of trait values in many populations is not homogenous but creates a mosaic of patches. This may lead to differences in selection on the patch level compared to selection on the population level. As an example we investigated the spatial distribution of nectar production and its effects on pollinator behaviour in a natural population of Echium vulgare. Nectar production per flower, number of flowers and total nectar production showed a hierarchy and spatial aggregation as expressed by Gini coefficients and significant Moran's I values. Plants in patches of high nectar production received significantly more pollinator visits and had a significant emanating effect on pollinator visits of neighbouring plants. The same was true for plants in patches with high number of flowers. To disentangle these effects a path analysis was applied, which suggested that the direct effect of nectar production per flower although present, seems to be small compared to the effect of the number of flowers. Nectar production per flower affected pollinator visits mainly indirectly by way of total nectar production, which includes the effect of number of flowers. Assuming a minor pollinator-mediated selection for number of flowers, pollinator-mediated selection for total nectar production equals that for nectar production per flower. If so, the observed spatial structure of nectar production and its emanating effect on pollinator behaviour is of importance for natural selection. Plants of low nectar production occurring close to patches of plants with high nectar production benefited from the enhanced pollinator service of their neighbours while saving costs of increased nectar production. Consequently, plants with low nectar production may have a selective advantage at patch level while plants with high nectar production may have a selective advantage at population level. Results presented stress the importance of small-scale patterns for ecological relationships and evolutionary change.     

The effects of a bumble bee nectar robber on plant reproductive success and pollinator behavior

Interactions between a plant species (Corydalis caseana), a bumble bee nectar robber (Bombus occidentalis), and a bumble bee pollinator (B. appositus) were studied. There were no significant differences between naturally robbed and unrobbed flowers in fruit set or mean seed set per fruit. Plots of C. caseana plants were subjected to treatments of robbing and no robbing using commercially available colonies of B. occidentalis. Robbers did not pollinate the flowers. Pollinator behavior was observed to determine (1) the number of bees attracted to each plot, (2) the number of inflorescences visited in a plot, (3) the number of flowers visited on each inflorescence, and (4) the distance flown between inflorescences. There were no significant differences in the number of inflorescences visited per bee or the number of flowers visited per inflorescence per bee when robbed and unrobbed treatments were compared. Of the parameters measured, only distance flown between inflorescences differed in the robbed and the unrobbed treatments. Bees flew significantly further between inflorescences in the robbed plots than in the unrobbed plots. The results indicate that the nectar robbers have no negative effect on fruit set or seed set in C. caseana and that they may cause increased pollen flow distances by changing the behavior of the pollinator.     

Flower and nectar attributes of pepper (Capsicum annuum L.) plants in relation to their attractiveness to honeybees (Apis mellifera L.)

Flower morphology, nectary structure, nectar traits and rates of honeybee foraging on pepper plants were studied. The nectary appears as swellings on the basal part of the ovary. The nectariferous cells are smaller and denser than the neighbouring parenchyma. Stomata are present in the nectary epidermis, but do not appear on the other parts of the ovary epidermis. Seven pepper breeding lines were grown near a bee yard in Rehovot. Five to six fold differences in nectar volume were found between the extreme genotypes. Nectar volumes were higher during noon and afternoon hours, as compared with morning hours. High correlation coefficients between nectar volume and sugar concentration were found. These were significant for the four high nectar yielding genotypes, ranging between r = 0.65 to r = 0.94. Male-fertile flowers produced more nectar and higher sugar concentration than sterile ones. Skewed distribution was observed in nectar volume of F₂ populations, but relatively low heritability values were calculated. Pepper nectar contains fructose and glucose only. The former occupies 52–82 % of the total sugar content. Pepper genotypes varied in frequency of honeybee visits and significant correlation between sugar quantity and number of honeybee visits per flower was evident. Fertile pepper flowers are not very attractive to honeybees and male-sterile flowers are even less so. The considerable variation in nectar characteristics can be exploited to increase attractiveness to honeybees, thus facilitating bee pollination in commercial production of F₁hybrid seeds and improve fruit quality.     

Nectar robbery by bees Xylocopa virginica and Apis mellifera contributes to the pollination of rabbiteye blueberry

Honey bees, Apis mellifera L., probe for nectar from robbery slits previously made by male carpenter bees, Xylocopa virginica (L.), at the flowers of rabbiteye blueberry, Vaccinium ashei Reade. This relationship between primary nectar robbers (carpenter bees) and secondary nectar thieves (honey bees) is poorly understood but seemingly unfavorable for V. ashei pollination. We designed two studies to measure the impact of nectar robbers on V. ashei pollination. First, counting the amount of pollen on stigmas (stigmatic pollen loading) showed that nectar robbers delivered fewer blueberry tetrads per stigma after single floral visits than did our benchmark pollinator, the southeastern blueberry bee, Habropoda laboriosa (F.), a recognized effective pollinator of blueberries. Increasing numbers of floral visits by carpenter bee and honey bee robbers yielded larger stigmatic loads. As few as three robbery visits were equivalent to one legitimate visit by a pollen-collecting H. laboriosa female. More than three robbery visits per flower slightly depressed stigmatic pollen loads. In our second study, a survey of 10 commercial blueberry farms demonstrated that corolla slitting by carpenter bees (i.e., robbery) has no appreciable affect on overall V. ashei fruit set. Our observations demonstrate male carpenter bees are benign or even potentially beneficial floral visitors of V ashei. Their robbery of blueberry flowers in the southeast may attract more honey bee pollinators to the crop.     

Bee visitation rates to cultivated sunflowers increase with the amount and accessibility of nectar sugars

Pollinators make foraging decisions based on numerous floral traits, including nectar and pollen rewards, and associated visual and olfactory cues. For insect‐pollinated crops, identifying and breeding for attractive floral traits may increase yields. In this study, we examined floral trait variation within cultivated sunflowers and its effects on bee foraging behaviours. Over 2 years, we planted different sunflower inbred lines, including male‐fertile and male‐sterile lines, and measured nectar volume, nectar sugar concentration and composition, and corolla length. During bloom, we recorded visits by both managed honey bees and wild bees. We then examined consistency in relative nectar production by comparing field results to those from a greenhouse experiment. Sunflower inbred lines varied significantly in all floral traits, including the amount and composition of nectar sugars, and in corolla length. Both wild bee and honey bee visits significantly increased with nectar sugar amount and decreased with corolla length, but appeared unaffected by nectar sugar composition. While wild bees made more visits to sunflowers providing pollen (male‐fertile), honey bees preferred plants without pollen (male‐sterile). Differences in nectar quantity among greenhouse‐grown sunflower lines were similar to those measured in the field, and bumble bees preferentially visited lines with more nectar in greenhouse observations. Our results show that sunflowers with greater quantities of nectar sugar and shorter corollas receive greater pollination services from both managed and wild bees. Selecting for these traits could thus increase sunflower crop yields and provide greater floral resources for bees.     

Sex differential nectar secretion in protandrous Alstroemeria aurea (Alstroemeriaceae): is production altered by pollen removal and receipt?

We examined diurnal and nocturnal nectar secretion across sexual stages in protandrous Alstroemeria aurea, a bumble bee-pollinated herb with long-lived flowers native to the southern Andes. We found the following patterns: (1) most nectar was produced diurnally and (2) three times more sugar was secreted during the male than female phase, not only because the male phase lasted longer but also because the rate of nectar production was higher. This 3:1 ratio in nectar production matched the ratio of the minimum number of bumble bee visits required on average to saturate male (pollen removal) vs. female (seed set) functions. Standing crop of nectar, on the other hand, did not differ greatly between male- and female- stage flowers left open to visitors, because the high-production male-phase flowers were visited more frequently than female- phase flowers. In an experiment concurrent with the repeated nectar sampling of individual flowers over their life-span, we removed pollen from anthers or deposited pollen on stigmas by hand. Neither treatment, designed to mimic effects of visits by Alstroemeria's native bumble bee pollinator, affected nectar production. The absence of plasticity in nectar secretion in relation to pollination events may reflect a low cost of nectar production, or may result from developmental constraints related to the evolution of the synchronous protandry that characterizes A. aurea.     

Effects of nectar theft by flower mites on hummingbird behavior and the reproductive success of their host plant, Moussonia deppeana (Gesneriaceae)

Hummingbird flower mites are transported in the nares of hummingbirds and may compete with them by "robbing" nectar secreted by the host plants. We have shown that Tropicoseius sp. flower mites consume almost half the nectar secreted by the long-lived, protandrous flowers of Moussonia deppeana (Gesneriaceae) pollinated by Lampornis amethystinus (Trochilidae). In this paper, we ask whether mimicking nectar consumption of flower mites alters some aspects of hummingbird foraging patterns, and, if so, how this affects host plant seed production. We observed hummingbirds foraging on (a) plants in which nectar was removed from the flowers and then filled with a sugar solution to half the volume of nectar simulating nectar consumption by flower mites, and (b) plants where nectar was removed and then filled with the sugar solution up to normal nectar volumes. Flower mites were excluded from both groups of plants to control for mite activity. Hummingbirds made fewer but longer visits to plants and revisited more the flowers with nectar removal than those without the treatment. We then conducted a pollination experiment on pistillate flowers using a stuffed L. amethystinus hummingbird to evaluate the effect of pollination intensity (number of bill insertions into one flower) on seed production. Flowers with more insertions produced significantly more seeds than those flowers that received fewer insertions. We conclude that the simulation of nectar consumption by hummingbird flower mites can influence the behavior of the pollinator, and this may positively affect seed production.     

Pollen carryover,nectar rewards,and pollinator behavior with special reference to Diervilla lonicera

Summary Pollen carryover was measured in three species of bumble bee pollinated plants by counting the numbers of foreign grains applied to the stigmas of a series of flowers by bumble bees. Deposition declined with the number of flowers visited in a roughly exponential fashion; most grains were deposited on the first few flowers, but some grains went much farther, the maximum carryover being 54 flowers. Variation in deposition was very high. In Diervilla lonicera, bees desposited significantly more grains on flowers which contained large amounts of nectar than on drained flowers. The implications are discussed in terms of plant strategies for optimizing pollination.     

Pollination ecology of Anthyllis vulneraria subsp. vulgaris (Fabaceae): nectar robbers as pollinators

This paper examines the hypothesis that nectar robbing can affect plant reproductive success either positively or negatively. To this end, I investigated various aspects of the pollination ecology of a population of the herb Anthyllis vulneraria subsp. vulgaris in northwest Spain over 5 yr. By observing floral visitors, I found that the most important pollinator species was the long-tongued bee Anthophora acervorum, which accounted for ～45% of recorded insect visits. However, just over 45% of visits were by the nectar-robbing bumble bees Bombus terrestris and B. jonellus. Although the incidence of robbing differed considerably over 5 yr of study, the frequency in every season was very high (66.4-76.5% of robbing) except for 1997 (0% robbing). Despite this high frequency of robbing, robbed flowers had a higher probability of setting fruit than nonrobbed flowers in all years of the study (mean: 82.0 vs. 51.0%; excluding 1997). This increased fruit set in robbed flowers is directly related to bumble bee behavior because the robbers' bodies came into contact with both the anthers and stigmas while robbing. Thus, the robbers effect pollination. These results suggest that the effect of nectar robbers on plant reproductive success is dependent both on the robbers' behavior and on flower/inflorescence structure. The importance of nectar-robbing bumble bees on the reproductive success of A. vulneraria and its yearly high frequency suggest that the relationship between robbers and this plant is part of a successful long-term mutualism.     

Unpredictability of nectar nicotine promotes outcrossing by hummingbirds in Nicotiana attenuata

Many plants use sophisticated strategies to maximize their reproductive success via outcrossing. Nicotiana attenuata flowers produce nectar with nicotine at concentrations that are repellent to hummingbirds, increasing the number of flowers visited per plant. In choice tests using native hummingbirds, we show that these important pollinators learn to tolerate high‐nicotine nectar but prefer low‐nicotine nectar, and show no signs of nicotine addiction. Nectar nicotine concentrations, unlike those of other vegetative tissues, are unpredictably variable among flowers, not only among populations, but also within populations, and even among flowers within an inflorescence. To evaluate whether variations in nectar nicotine concentrations increase outcrossing, polymorphic microsatellite markers, optimized to evaluate paternity in native N. attenuata populations, were used to compare outcrossing in plants silenced for expression of a biosynthetic gene for nicotine production (Napmt1/2) and in control empty vector plants, which were antherectomized and transplanted into native populations. When only exposed to hummingbird pollinators, seeds produced by flowers with nicotine in their nectar had a greater number of genetically different sires, compared to seeds from nicotine‐free flowers. As the variation in nectar nicotine levels among flowers in an inflorescence decreased in N. attenuata plants silenced in various combinations of three Dicer‐like (DCL) proteins, small RNAs are probably involved in the unpredictable variation in nectar nicotine levels within a plant.     

Variability in nectar production and standing crop, and their relation to pollinator visits in a Mediterranean shrub

Nectar standing crops in flowers within an individual plant are often highly variable. This variability may be a by-product of the foraging activity of insect pollinators. Alternatively, plants may be selected to produce highly variable rewards to reduce consecutive visitation by risk-averse pollinators, thus diminishing within-plant pollen transfer. This study evaluated the roles of pollinator control vs. plant control over nectar variability in the bee-pollinated shrub Rosmarinus officinalis L. (Lamiaceae). We sampled nectar production, standing crop and pollinator visits in three shrubs of one population over 17 days during one blooming season. Nectar production rates were highly variable (CV = 1.48), and increased after rainy days. Nectar standing crops were even more variable (CV = 2.16), decreased with increasing temperatures, and increased with time since the last rain. Pollinator visit rates decreased with variability in nectar standing crops, increased with flower number per shrub, and were unaffected by variability in nectar production rates. Repeated sampling of marked flowers revealed no correlation between their nectar standing crops and production rates. These findings support the role of reward variance in reducing pollinator visits, but suggest that plants are not in complete control of this variability. Rather, plant-generated variability can be modified by intensive foraging activity of pollinators. Such pollinator control over nectar variability is likely to reduce the selective advantage of plant-generated reward variation. Handling Editor: Neal Williams.     

Novel nectar robbing negatively affects reproduction in Digitalis purpurea

With many plant–pollinator interactions undergoing change as species’ distributions shift, we require a better understanding of how the addition of new interacting partners can affect plant reproduction. One such group of floral visitors, nectar robbers, can deplete plants of nectar rewards without contributing to pollination. The addition of nectar robbing to the floral visitor assemblage could therefore have costs to the plant´s reproductive output. We focus on a recent plant colonist, Digitalis purpurea, a plant that in its native range is rarely robbed, but experiences intense nectar robbing in areas it has been introduced to. Here, we test the costs to reproduction following experimental nectar robbing. To identify any changes in the behavior of the principal pollinators in response to nectar robbing, we measured visitation rates, visit duration, proportion of flowers visited, and rate of rejection of inflorescences. To find the effects of robbing on fitness, we used proxies for female and male components of reproductive output, by measuring the seeds produced per fruit and the pollen export, respectively. Nectar robbing significantly reduced the rate of visitation and lengths of visits by bumblebees. Additionally, bumblebees visited a lower proportion of flowers on an inflorescence that had robbed flowers. We found that flowers in the robbed treatment produced significantly fewer seeds per fruit on average but did not export fewer pollen grains. Our finding that robbing leads to reduced seed production could be due to fewer and shorter visits to flowers leading to less effective pollination. We discuss the potential consequences of new pollinator environments, such as exposure to nectar robbing, for plant reproduction.     

Behavioural differences between male and female carpenter bees in nectar robbing and its effect on reproductive success in Glechoma longituba (Lamiaceae)

Male and female nectar robbers may show significantly different behaviour on host plants and thus have different impacts on reproductive fitness of the plants. A 4-year study in natural populations of Glechoma longituba has shown that male carpenter bees (Xylocopa sinensis) are responsible for most of the nectar robbing from these flowers, while female bees account for little nectar robbing, demonstrating distinct behavioural differentiation between male and female bees in visiting flowers. The smaller male bee spends less time visiting a single flower than the larger female bee, consequently, the male bee is capable of visiting more flowers per unit time and has a higher foraging efficiency. Moreover, the robbing behaviour of female carpenter bees is more destructive and affects flower structures (ovules and nectaries) and floral life-span more than that of the male bee. According to the energy trade-off hypothesis, the net energy gain for male bees during nectar robbing greatly surpasses energy payout (17.72 versus 2.43 J), while the female bee net energy gain is barely adequate to meet energy payout per unit time (3.78 versus 2.39 J). The differences in net energy gain for male and female bees per unit time in nectar robbing are the likely cause of observed behavioural differences between the sexes. The differences in food resource preference between male and female bees constitute an optimal resource allocation pattern that enables the visitors to utilise floral resources more efficiently.