The superiority of bumblebees to honeybees as pollinators: insect visits to raspberry flowers

Abstract.

• 1 The behaviour and activity patterns of Apis mellifera and of five species of Bombus were analysed in relation to climatic variables and nectar quality on three varieties of unsprayed cultivated raspberry (Rubus idaeus) in eastern Scotland.
• 2 Stages of floral morphology and reward were similar for the three varieties: young flowers offered both nectar and pollen, but medium and old flowers offered nectar only, in diminishing quantities.
• 3 A wide range of insects visited raspberry flowers, but bees were dominant, bumblebees being responsible for about 60% of all visits and honeybees making up most of the remaining percentage. All bees had substantial pollen deposited on their bodies during visits, though few specifically collected it.
• 4 Bombus spp. were found to favour young (receptive) flowers strongly, especially early in the morning when pollen was most abundant: whilst Apis visited unselectively. Bumblebees also foraged over substantially longer periods of the day, and in poorer weather, some being present at most times of observation; and they foraged more quickly in terms of flower visits per minute.
• 5 Bombus carried more pollen on their bodies than Apis, and also deposited more pollen on raspberry stigmas, with B.lapidarius and B.terrestris being particularly effective and also being the most abundant species. All bumblebees also foraged over a longer range, moving between canes and rows more frequently than did honeybees.
• 6 Bumblebees are therefore likely to be substantially more important as pollinators of raspberries than are honeybees, especially as raspberries though moderately self-fertile may exhibit metaxenia. Reasons why Bombus may be the preferred pollinator in most sites of raspberry cultivation are discussed, together with implications for present and future growers.

     

The presence of aggressive ants is associated with fewer insect visits to and altered microbe communities in coffee flowers

The process of dispersal can shape ecological communities, but its influence is thought to be small compared to the effects of environmental variation or direct species interactions, particularly for microbial communities. Ants can influence movement patterns of insects and the microbes they vector, potentially affecting microbial establishment on plants, including in agroecosystems. Here, we examine how the presence of aggressive ants, which can influence floral visitation by bees and other pollinators, shapes the community composition of bacteria and fungi on coffee flowers in farms that differ in shade management intensity. We hypothesized that the presence of aggressive ants should reduce the frequency and diversity of floral visitors. Finally, we predicted that the effects of ants should be stronger in the low-shade farm, which has a less diverse community of floral visitors. We sampled microbial communities from nectar and pistils of coffee flowers near and far from nests of the aggressive ant Azteca sericeasur across two farms that vary in shade management and diversity of floral visitors. Bacterial and fungal community composition was characterized using Illumina sequencing of the 16S and ITS regions of the rRNA gene. Consistent with our expectation, Azteca presence was associated with a decrease in the number and diversity of visitors, visit duration and number of flowers visited. Azteca presence influenced microbial communities, but effects differed between farms. Azteca nests were associated with higher bacterial diversity in both farms, but the difference between flowers on trees with and without Azteca was greater in the high-shade farm. Azteca nests were associated with higher fungal diversity in the high-shade farm, but not the low-shade farm. In addition, the presence of ants was strongly associated with species composition of fungi and bacteria in flowers, but differentiation between ant and no-ant communities was greater in the low-shade farm. Specific operational taxonomic units (OTUs) were differentially associated with the presence of ants. We conclude that indirect interactions that influence dispersal may have large effects on microbial community composition, particularly in ephemeral microbial communities.     

A trade-off between the frequency and duration of bumblebee visits to flowers

Pollinator behavior influences plant reproduction in many ways. A traditional measure of pollination, the number of visits received, may be a poor predictor of plant reproductive success, particularly when there are trade-offs between visit quantity and components of visit quality. For example, the duration of pollinator visits may be negatively correlated with the number of visits received by a flower. We tested for a trade-off between the number of bumblebee visits and the duration of those visits in an experimental population of snapdragons (Antirrhinummajus: Scrophulariaceae). The duration of a bumblebee visit to a flower increased significantly with the time interval since the flower had last been visited. Over the lifetime of a flower the correlation between the total number and average duration of visits received by a flower was weakly negative. However, at the whole-plant level the correlation was positive: plants whose flowers received more visits also received visits of longer duration. Factors affecting the relationship between quantity and duration of pollinator visits to flowers also were investigated. Two factors weakened the negative dependence of average visit duration on number of visits received by individual snapdragon flowers: (1) the correlation between the total number of visits to a flower and the average interval between visits was only −0.53, as visits to individual flowers were not very evenly spaced over time, and (2) newly opened flowers received fewer and shorter visits than older flowers. Comparing whole plants, nectar production per flower varied dramatically across individuals, a probable explanation for the positive correlation between visit number and average duration per flower observed at the plant level. The potential for a trade-off between these two components of pollinator service exists when visit duration depends on reward quantity; whether the trade-off is realized will depend on variation in nectar production and on whether pollinators forage systematically. Received: 3 October 1997 / Accepted: 16 June 1998     

Interaction between flowers, ants and pollinators: additional evidence for floral repellence against ants

It has been commonly suggested that ants negatively affect plant pollination, particularly in the tropics. We studied ant–flower–pollinator interactions in a lowland rainforest in Borneo. Frequency and duration of pollinator visits were compared between flowers attended by ants and flowers from which ants were excluded. In all four plant species studied, the activity of ants decreased the rate and/or duration of the pollinators’ floral visits. For this and other reasons it is expected that plants repel ants from flowers during anthesis. We tested this prediction for a different set of plant species in which we observed the behaviour of Dolichoderus thoracicus ants when encountering flowers. In eight out of 18 plant species studied, ants showed a significantly higher rejection rate when they encountered flowers than when they encountered controls. Our results are thus consistent with the hypothesis that ants may negatively affect plant fitness by reduced intensity of pollinator visits and that ants are repelled from flowers of many tropical plant species, although this repellence is clearly not ubiquitous.     

Dynamics of forager arrivals and nectar renewal in flowers of Anchusa strigosa

Summary Long-tongued Anthophora spp. bees collecting nectar from flowers of Anchusa strigosa (Boraginaceae) exhibit systematic foraging. Successive forager arrivals at individual flowers are not independent, and the time elapsed between successive arrivals at a particular flower is distributed more uniformly than be expected on the basis of a random arrival process. Distributions of inter-arrival time for individual flowers show standard deviation/mean ratios of 0.44–0.79, a range which is consistent with results obtained for two other plant-pollinator systems. The rate at which nectar is renewed between successive forager arrivals is independent of the amount of nectar in the flower, and the renewal process is strongly linear. Practical and theoretical implications of these results are discussed.     

The colour of flowers in spectrally variable illumination and insect pollinator vision

The spectral reflectance of differently coloured Australian native plant flowers and foliage was measured and plotted in a colour triangle to represent the colour space of the honeybee. Spectral variations in illumination are shown to significantly change plant colours for bee vision without colour constancy. A model of chromatic adaptation based upon the von Kries coefficient law shows a reduction in plant colour shift, with the degree of correction depending upon position in colour space. A set of artificial reflectances is used to map relative colour shift caused by spectrally variable illumination for the entire colour space of the honeybee. The rarity of some flower colours in nature shows a correlation to a larger colour shift for these colours when illuminated by spectrally variable radiation. The model of chromatic adaptation is applied to illuminations used in a behavioural study on honeybee colour constancy by Neumeyer 1981. Surface colours used by Neumeyer are plotted in colour space for the various illuminations. The results show that an illumination-dependent colour shift correlates to a decrease in the frequency of bees correctly choosing a colour to which it was trained. Accepted: 23 February 1998     

Diversity,abundance and foraging behaviour of insect pollinators in Radish (Raphanus raphanistrum subsp. sativus L.)

Insect pollinators and flowering plants have co-evolved together for millions of years, developing a wide array of mutualistic interactions. Several cultivated plant species depend exclusively on insect-mediated pollination services for their subsistence, reproduction and survival. Radish, Raphanus raphanistrum subsp. sativus, a popular edible root vegetable (Family: Brassicaceae) completely depends on pollinators as self-pollination is totally absent due to sporophytic self-incompatibility. The present study was conducted at the experimental fields of the ICAR-Indian Institute of Horticultural Research, Bengaluru, to understand the diversity and foraging behaviour of insect pollinators in R. sativus. Continuous survey during the flowering period revealed that a total of fifteen insect species that belong to three insect orders (Hymenoptera, Diptera and Lepidoptera) were found to visit R. sativus flowers. The most abundant species was Apis florea L., followed by Apis dorsata F., Tetragonula iridipennis (Smith), Apis cerana F., Lassioglossum sp. and Ischiodon scutellaris F. Floral visitors differed significantly in terms of their visitation frequency, with A. florea (0.76 ± 0.22 visits/flower/ min) being the most frequent visitor followed by A. dorsata (0.66 ± 0.21 visits/flower/ min) and T. iridipennis (0.59 ± 0.14 visits/flower/30 min). The maximum visitation rate was recorded for A. dorsata (7.8 ± 1.40 flowers visited/min) followed by A. cerana (6 ± 1.15). The maximum stay time on individual flower was recorded for Lassioglossum sp. (48.2 ± 3.67 sec) followed by T. iridipennis (44.2 ± 4.24 sec). The maximum numbers of pollen grains were deposited by A. dorsata (206.70 ± 56.45) followed by A. cerana (151.20 ± 31.84). All the floral visitors except A. florea made contact with stigma on every single visit (100 %). While, only 53 per cent of total visitation by the Apis florea led to stigma contact. Further, nectar robbing behaviour was majorly exhibited by A. florea and Delta conoideum compared to all other insect pollinators observed. Based on the overall pollinator effectiveness per day A. dorsata was found to be the most effective single pollinator species.     

Bouncy versus idles: On the different role of pollinators in the generalist Gentiana lutea L.

Generalist flowers are visited by a broad variety of insects that function as pollinators, occasional visitors and as pollen and/or nectar robbers. Moreover, among legitimate pollinators the pollination efficiency can be different. Nectar greatly affects visitor behaviour and fidelity to a certain species, influencing plant reproductive effort. In this study we have investigated a generalist system (Gentiana lutea L.), examining the role of flower visitors and quantifying the contribution of each pollinating taxon in three natural populations. In order to verify the level of generalization, we introduce an index of Pollinator Performance (PoP), based on insect visitation rate and cross-pollen transport efficiency. Our results confirm the high degree of pollinator-generalization of the study species. Nevertheless, flower visitors show various degree of pollinating performance, mainly defined by their sedentary versus dynamic behaviour. Sedentary insects enhance geitonogamous pollen transfer, which results in reduced seed set and pollen limitation. In particular, an unusual sluggish behaviour was observed in bumblebees feeding on nectar. The hexose-rich abundant nectar offered by G. lutea flowers is remarkably rich in proline and β-alanine amino acids: this composition presumably influences feeding choice and insect dynamism, likely exerting a narcotic effect on pollinators. The consequences on plant fitness are discussed in an evolutionary perspective.     

Use of visual and olfactory cues of flowers of two brassicaceous species by insect pollinators

1. Pollinating insects exploit visual and olfactory cues associated with flower traits indicative of flower location and reward quality. Pollination can induce changes in these flower-associated cues, thereby influencing the behaviour of flower visitors. 2. This study investigated the main cues exploited by the syrphid fly Episyrphus balteatus and the butterfly Pieris brassicae when visiting flowers of Brassica nigra and Raphanus sativus plants. Whether pollen is used as a cue and whether pollination-induced changes affect flower volatile emission and the behavioural responses of the two pollinator species were also studied. 3. Pollinator preference was investigated by offering visual and olfactory cues individually as well as simultaneously in two-choice bioassays. Plant treatments included emasculation, hand-pollination and untreated control plants. The composition of flower volatiles from pollinated and unpollinated control plants was analysed by gas chromatography-mass spectrometry. 4. Both pollinators exhibited a strong bias for visual cues over olfactory cues. Neither pollinator used pollen as a cue. However, E. balteatus discriminated between newly opened and long-open flowers at short distance only when pollen was available. Flower visits by pollinators were influenced by pollination-induced changes in B. nigra but not R. sativus flowers. Pieris brassicae only responded to pollination-induced changes when visual and olfactory cues were offered simultaneously. The blend of volatiles emitted by B. nigra, but not R. sativus inflorescences was affected by pollination. 5. Collectively, the findings of this study show that different pollinators exploit different visual and olfactory traits when searching for flowers of two brassicaceous plant species.     

Effects of inflorescence size on visits from pollinators and seed set of Corydalis ambigua (Papaveraceae)

Female reproductive success (seed set) of a spring ephemeral plant, Corydalis ambigua Cham. et schlecht (Papaveraceae) was investigated in relation to inflorescence size and foraging behavior (frequency and duration of visitations) by pollinators (namely, overwintered queens of Bombus hypocrita sapporensis) by detailed daily observations of a natural population. Pollination experiments indicated that C. ambigua is self-incompatible and that seed set was significantly affected by the behavior of the pollinating queens. Plants with larger inflorescences were visited more often than those with fewer flowers. Fecundity also increased with increasing size of inflorescences. Visitation time (duration of foraging) rather than the frequency of visitations (number of visits) was critical for higher fecundity. Seed production was strongly enhanced by a few long visits (of more than 60 s), and seemed to be independent of large numbers of short visits (of less than 60 s). Hence, plants with larger inflorescences, which provide a conspicuous signal to pollinators and offer greater rewards in terms of nectar, received longer visits by B. hypocrita sapporensis queens and those plants exhibited higher fecundity.     

Floral color change and the attraction of insect pollinators in lungwort (Pulmonaria collina)

We studied the effect of floral color change on long- and short-distance attraction of insect pollinators to the herb lungwort, Pulmonaria collina. Lungwort flowers change color with age from red to blue. Young red flowers had a significantly greater pollen and nectar reward and were significantly more often unpollinated than old blue ones. Red and blue flowers both influenced long-distance attractiveness of plants, defined as the number of insect approaches towards an individual plant. After reaching a plant, flower visitors preferred to visit young red flowers. Therefore, short-distance attractiveness, defined as the number of flowers visited successively on an individual plant, was influenced mainly by the number of young red flowers. The co-occurrence of the change in reproductive ability, in amount of reward, and in flower color enabled lungwort plants to direct pollinators to reproductive, highly rewarding red flowers. The data suggest that by maintaining changed flowers lungwort plants can increase their long-distance attraction and simultaneously enhance the probability of flower visits to pre-changed flowers. Thus, we propose floral color change as a mechanism that can increase the efficiency of pollen transfer to enhance plant fitness. Received: 2 November 1998 / Accepted: 14 July 1999     

Nectar and pollen sources for honeybees in Kafrelsheikh province of northern Egypt

This study was conducted at the apiary of the Beekeeping Research Section at the Sakha Agricultural Research Station, ARC, Kafrelsheikh, and other apiaries in Kafrelsheikh province, during two successive years 2015 and 2016. The study aimed to survey nectar and pollen floral resources in Kafrelsheikh province. Ninty seven plant species belonging to 33 families were recorded as nectar sources, and 82 plant species belonging to 36 families were recorded as pollen sources during the whole year. The largest amount of monthly trapped pollen was obtained during May followed by August. It can be concluded that, beekeepers in Kafrelsheikh province can harvest good honey yield at the end of blooming seasons of citrus (Citrus spp.) during March and April, Egyptian clover (Trifolium alexandrinum L.) during May and June, loofah (Luffa aegyptiaca Mill.) during June to October, cotton (Gossypium spp.) during July and August, and banana (Musa spp.) during August and September. They also, could be trapping pollen loads collected from faba bean (Vicia faba L.) and flax (Linum usitatissimum L.) during January to March, date palm (Phoenix dactylifera L.) during March and April, Egyptian clover during May and June, summer seed watermelon (Citrullus lanatus var. colothynthoides L.) during June and July, loofah and maize (Zea mays L.) during June to November.     

Bumblebee visits to Impatiens spp.: pattern and efficiency

Summary Three Japanese species of Impatiens, which secrete nectar continuously in long spurs, were visited by Bombus diversus workers consecutively throughout the day. B. diversus workers showed characteristic patterns of behavior in flower use, flower choice, and patch departure. (1) Bumblebees stayed longer on a flower which had been unvisited for a while than on a flower which had been visited recently. (2) Bumblebees preferred visiting flowers which had been unvisited for a while to visiting those which had been visited recently, and to visiting those which had been unvisited for a long period. (3) Bumblebees had a higher probability of leaving a patch after they had stayed on a flower for a short period than after they had stayed for a longer period. The bumblebees appeared to perceive both remotely and proximately chemical cues deposited by other foraging individuals, which indicated nectar rewards in a flower, and thus obtained a higher nectar intake than the mean amount of nectar left in a flower.     

花部重金属积累对植物-传粉昆虫互惠关系影响的研究进展

传粉昆虫急剧下降是当前全球生态学家最为关注的热点问题之一,全球气候变化、土地利用改变、生境破碎化以及工农业生产带来的有害物质如杀虫剂等均有重要影响。相比而言,土壤重金属污染对传粉昆虫的潜在影响缺乏深入评价。土壤重金属会进入植物组织如花部,在传粉过程中传递到传粉者体内,并通过影响传粉者行为改变植物雌雄适合度,全面梳理相关研究进展和存在的问题,有助于提升对重金属污染带来的生态后果的全面认识,了解动植物相互关系对环境变化的响应。本文结合前期的研究成果,从重金属在植物花部中的积累模式、花部重金属积累对植物雌雄适合度的影响、经花部报酬介导的重金属积累对蜂类传粉者传粉行为、生活史关键环节的影响等进行综述,以期为理解花部重金属积累对植物与传粉昆虫互惠关系的影响,进而拓展关于土壤重金属污染与传粉昆虫数量减少之间的内在联系的科学认识。     

A meta-analysis of physiological and behavioral responses of parasitoid wasps to flowers of individual plant species

This meta-analysis summarizes published information on the use of various plant species by parasitoid wasps. Trials measured either a physiological response such as wasp longevity when supplied with flowers from a single plant species, or a behavioral response like attraction of wasps to field plantings. The log response ratio was the effect size used to standardize estimates and meta-analyses were conducted to make overall estimates for plant species included in multiple trials. Physiological response trials have been conducted on 126 different plant species and behavioral response trials tested 104 plant species. The log response ratio of different plant species ranged from 0 to 2.7 for longevity, and up to 4.1 for attraction. The longevity response ratio estimate is equivalent to a nearly 15-fold increase in the ratio of days longevity with the flower to days longevity in the starvation control. Wasp longevity increased the most with plants in the Polygonaceae, Apiaceae, Brassicaceae, Boraginaceae, Solanaceae, and Rosaceae. Plant species in the Onagraceae, Caryophyllaceae, Lamiaceae, Scrophulariaceae, Asteraceae, and Fabaceae tended to slightly increase wasp life spans, but their effect was lower and less consistent among species. There were a number of families which did not increase wasp longevity over the controls at all such as Chenopodiaceae. This review can help identify plant species which have been proven to supply nectar for parasitoids for potential use in a conservation biological control program, but plant selection should not be limited to the small list of species that were included in these studies.     

Some pollinators are more equal than others: Factors influencing pollen loads and seed set capacity of two actively and passively pollinating fig wasps

The nursery pollination system of fig trees (Ficus) results in the plants providing resources for pollinator fig wasp larvae as part of their male reproductive investment, with selection determining relative investment into pollinating wasps and the pollen they carry. The small size of Ficus pollen suggests that the quantities of pollen transported by individual wasps often limits male reproductive success. We assessed variation in fig wasp pollen loads and its influence on seed production in actively pollinated (Ficus montana) and passively pollinated (Ficus carica) dioecious fig trees.The ratios of number of male flowers on number of female flowers in a glasshouse-maintained F. montana population were highly variable. When fig wasps were introduced into receptive female figs, the resulting seed numbers were strongly linked to the numbers of pollinators that had been seeking access to pollen, relative to the number of anthers in their natal figs. In F. carica estimates of the amounts of pollen produced per fig and the quantities of pollen carried by emerging fig wasps suggest that less than 10% of the pollen is transported. Pollinators of F. carica that emerged earlier from figs carried more pollen, and also generated more seeds when introduced into receptive female figs.We show here that all pollinators are not equally valuable and producing more pollinators is not necessarily a good option in terms of Ficus male fitness. Previous results on F. montana figs showed that only around half of the flowers where pollinators lay eggs produced adult offspring. The amount of pollen collected by young female fig wasps may be a major determinant of their reproductive success.     

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.     

GM risk assessment: Pollen carriage from Brassica napus to B. rapa varies widely between pollinators

Characterizing insect pollen carriage between closely related plant species is especially challenging where source species possess morphologically identical pollen and share many pollinators in common. Here, we use an SNP-based assay using the plant DNA barcoding locus matK to characterize pollen carriage between cultivated Brassica napus and wild Brassica rapa in three sites across southern England. The assay differentiated B. napus and B. rapa pollen carried by honey bees (Apis melifera), bumblebees (Bombus spp.), mining bees (Andrena spp.) and hoverflies (Syrphidae) captured on B. napus plants 1–2 m from wild B. rapa, and on B. rapa plants at various distances from the crop . Apis individuals foraging on B. rapa and carrying B. napus pollen were rarely found beyond 10 m from the crop. However, Bombus and Andrena individuals captured on B. rapa occasionally carried crop pollen up to 300 m from the source field. Hoverflies carried less pollen overall but featured high proportions of B. napus pollen even at the most distant capture points. We predict that different pollinator species will evoke markedly different patterns of interspecific hybrid formation. We conclude that more exhaustive surveys of this kind will help parameterize future mechanistic models to predict the distribution of hybrids between Genetically Modified B. napus and B. rapa on a landscape scale.     

Nectar replenishment and pollen receipt interact in their effects on seed production of Penstemon roseus

Resource supply and pollen delivery are often thought to equally limit seed production in animal-pollinated plants. At equilibrium, plants should show no response to experimental pollen supplementation because resources limit seed set above the current level of pollen attraction, while experimental reduction in pollen deposition below the equilibrium level would reduce seed set. The predicted equilibrium may be disrupted, however, if plants expend additional energy to replenish removed nectar. We investigated the combined effects of nectar removal and pollen delivery on female reproductive success of Penstemon roseus (Plantaginaceae), a hummingbird-pollinated plant that replenishes removed nectar. We first documented that the frequency of experimental nectar removal was correlated with total nectar secretion; and increased frequency of nectar removal resulted in increased female reproductive costs to the plant. Trade-offs between investing resources in nectar and investing resources in seeds were then investigated in two contrasting natural populations by removing nectar from flowers at increasing frequencies while simultaneously hand-pollinating flowers with increasing amounts of pollen. Seed set was lowest at low levels of pollen deposition, highest at medium-sized pollen loads, and intermediate when pollen loads were highest. At both sites, the frequency of nectar removal and pollen deposition had an interactive effect on seed production, in that intermediate levels of nectar removal result in the absolute highest seed set, but only at intermediate pollen loads. At high pollen loads, seed set was higher following little to no nectar removal, and at low pollen loads, all rates of nectar removal affected fecundity equally. Seed mass responded to nectar removal and pollination differently than did seed set. High levels of nectar removal and pollen delivery both lowered seed mass, with little interaction between main effects. Our findings are among the first to demonstrate that nectar replenishment costs and pollination intensity jointly affect seed production. This conflict between nectar replenishment costs and pollen-limiting factors results in trade-offs between pollinator attraction and seed provisioning. Thus, resource allocation towards nectar production should more often be considered in future studies of pollen limitation.