Western honey bee (Apis mellifera L.) attraction to commercial pollen substitutes and wildflower pollen in vitro

Many beekeepers feed their western honey bee (Apis mellifera) colonies artificial pollen substitutes to provide colonies with adequate nutrition during times of limited pollen quantity or quality. We provided caged worker bees commercially available pollen substitutes (AP23, MegaBee, UltraBee) and wildflower pollen in a choice-test to determine their relative attraction to/preference for the diets. We measured diet consumption by honey bees and observed honey bee behaviour to evaluate bee preferences for certain diets. The bees interacted with and consumed more wildflower pollen than they did any of the commercially available pollen substitutes. Our data suggest that bees have a strong preference for wildflower pollen over commercially available pollen substitutes.     

Nutritional effects of supplementary diets on brood development,biological activities and honey production of Apis mellifera L

The present research work was conducted to assess the impact of nutrient-enriched diet on the physiological activities and subsequently honey yield. Eighteen colonies of Apis mellifera L. were selected from Dera Ismail Khan region, KPK, Pakistan, during the winter and summer seasons, 2019–2020. Five pollen supplement diets were prepared and provided to screen out the palatable one to be fed as pollen alternative nutrition to bee bread. Results of diet consumption regarding mean data for consumption rate displayed that soybean flour enriched artificial diet was maximally consumed (74.34 g) by honey bees per week. Minimum consumption was observed for grinded groundnut enriched diet (64.62 g) which was relatively lesser than the other tested artificial diets. Results of area of worker brood disclosed that soybean flour fortified diet (1489.27 cm²/colony) statistically noteworthy than the other artificial diets whereas control (463.51 cm²/colony) was least effective. Highest bee strength (10.00 bee frames/colony) was noted in the bee colonies fed with soybean flour fortified diet, date paste (8.0 bee frames/colony) was the next effective one, among the tested pollen replacement diets whereas relatively least (5 bee frames/colony) was noticed in case of grinded groundnut. Highest body weight (12.41 g) of neonate bees was noted in case of soybean enriched diet while lowermost (5.31 g) was noted in control bees. Results of wax cell built up and foraging efficiency were also superior in artificial diets than respective control bees. Hence, artificial diets especially soybean-enriched pollen alternative diet can boost up the physiology of honey bee leading to increased honey yield and profit.     

Lipid-enhanced pollen and lipid-reduced flour diets and their effect on the longevity of honey bees (Apis mellifera L.)

Abstract  As eucalypt pollens contain low concentrations of lipid, enhancing pollen diets with fatty acids was hypothesised to improve honey bee longevity. Different concentrations of linoleic and oleic acid added to eucalypt pollen were trialled in small cages containing approximately 1400 bees each. Bees fed diets of redgum ( Corymbia calophylla (Lindl.) Hill & Johnson, formerly Eucalyptus calophylla ) pollen had the lowest mortality of 22 diets tested for 6 weeks and had life spans (50%) greater than 42 days. Linoleic acid mixed with a redgum diet in concentrations >6% corresponded to life spans of 24–25 days. Bee longevity appeared to be more sensitive to oleic acid as life spans decreased to 15–21 days when diets had concentrations >2%. The life spans of bees fed soya bean flour were 26 days on low (0.6% lipid) fat, 19 days on defatted and 20 days on full-fat diets. Bees fed lupin flour had a life span of 23 days. Adding redgum pollen to lupin flour caused increased mortality, but addition of pollen to soya bean flour was beneficial. Thus, beekeepers who choose to utilise soya bean or lupin flours as protein substitutes to pollen will have bees with reduced longevity. Bees fed redgum pollen that had been dried, crushed, irradiated and hermetically stored in a cool room for several years had similar longevity to bees fed fresh-collected and frozen redgum pollen.     

Colony pollen reserves affect body size,sperm production and sexual development in males of the stingless bee <Emphasis Type="Italic">Melipona beecheii</Emphasis>

The production of male sexual offspring by social insect colonies is often strongly seasonal or resource-dependent. In stingless bees, males are produced in smaller numbers under conditions of low colony food reserves; whether such males are negatively affected in traits related to reproductive success is not known. We compared body size, sperm production and sexual maturity in Melipona beecheii males reared with experimentally supplemented or reduced pollen reserves, but with otherwise equal numbers of workers and equal quantities of honey reserves. We also studied the same traits in males collected from non-manipulated colonies with pollen reserves intermediate between the supplemented or reduced groups but with more workers and honey reserves. Males reared under experimentally reduced pollen reserves had significantly smaller bodies and lower sperm counts compared to those reared in colonies with experimentally supplemented pollen reserves. There was also a significantly positive relationship between the number of sperm and body size in males across all colony treatments. The maximum number of sperm in seminal vesicles was recorded 2 days later in males from colonies with reduced pollen compared to males from colonies with supplementary pollen. Males from non-manipulated colonies were intermediate in size, sperm count and speed of maturation. Our study documents for the first time the existence of large size variation in males of stingless bees that is related with the amount of pollen reserves in their natal colony. We conclude that a colony’s pollen reserves have a major impact on male body size, sperm production and speed of sexual maturity in this stingless bee, which may be the case in other social insects. Stingless bees are a good model system to study the balance between colony-level selection and individual-level selection on male sexually selected traits such as body size.     

Influence of pollen feeding and physiological condition on pesticide sensitivity of the honey bee Apis mellifera carnica

Summary In two consecutive years heavy bee mortality at end April/early May followed the use of pesticides classed as harmless for bees along road verges. It was thought that old weak winter bees had succumbed to a preparation otherwise innocuous. Extensive tests to reveal any links between the bees' physiological condition and pesticide sensitivity involved 6 hormone herbicides, 11 fungicides and 2 insecticides, all approved harmless for bees and functioning on them wholly or mainly as stomach poisons. As a rule bee sensitivity was measured as LD 50 per os, in smaller tests as percentage mortality. Amount and quality of pollen ingested in the first days of life affected the pesticide sensitivity of young and older bees. Bees fed adequate high quality pollen are less sensitive than counterparts fed inadequate or inferior pollen or pollen substitute; such differences persisted if the LD 50 was calculated for the same body weight. Pesticides containing manganese are an exception. To these, bees fed inadequate pollen are no more or even less sensitive than comparable well-fed bees. Pesticide sensitivity decreases generally from early to late summer. Quality of pollen available for larvae has no effect on poison sensitivity of imagines. Food supply conditions however exert a clear influence: tested with the same pesticides, hive bees from colonies having had a rich early food supply, and young bees bred then, are less sensitive than their counterparts having had moderate or no early food supply. Poison sensitivity of summer bees increases with age; most sensitive are old winter bees which had practiced broodcare in early spring.Inadequate pollen intake can be regarded as causing protein deficiency. Investigation of this in mammals and man indicate that the higher poison sensitivity in bees results from inhibition of the enzymatic decomposition of pesticides. For practical bee protection it is important that all organic fungicides tested are effectively harmless. Hormone hebicides can be ranked as practically harmless even for bees inadequately protein-fed, as long as the approved concentrations are observed. Our tests raised doubts however about the registration as harmless for bees of insecticides based on Endosulfan and Phosalon. Of interest in practice and for the official testing of pesticides are also the high pesticide sensitivity of old winter bees, the decrease in sensitivity of bees on a stable feed from early to late summer, and the sensitivity-reducing influence of pollen-rich food supply promoting development.It is important ecologically that pollens of different plant species vary in nutrient quality for the honey bee: there are perfectly worthless (conifers), poor-to-medium, and highly effective pollen types. As shown in this paper, these differences are relevant not only for the development of the physiological condition and breeding potential of the bee, but also for pesticide sensitivity. That bees gather worthless and poor-quality, sometimes even poisonous, pollen (some Ranunculus sp.) is evidently due to the phagostimulant present in all pollen types.     

Asynchrony between solitary bee emergence and flower availability reduces flower visitation rate and may affect offspring size

Climate change can disrupt plant-pollinator interactions when shifts in the timing of pollinator activity and flowering occur unequally (i.e., phenological asynchrony). Phenological asynchrony between spring-emerging solitary bees and spring-flowering plants may cause bees to experience food deprivation that can affect their reproductive success. However, the mechanisms underlying the effects of food deprivation on solitary bee reproduction remain unknown. We investigated 1) whether food deprivation caused by phenological asynchrony affects solitary bee reproduction by influencing female lifespan and/or visitation to flowers, and 2) the relationship between the magnitude of asynchrony and bee responses. We simulated phenological asynchrony by depriving emerged female Osmia cornifrons (a spring-active solitary bee species) of nectar and pollen for 0 to 16 days. Following asynchrony treatments, we used flight cages to monitor 1) post-treatment female lifespan, 2) flower visitation, and 3) reproduction (i.e., total offspring, offspring weight, sex ratio). We found that post-treatment female lifespan was not affected by phenological asynchrony treatments, but that flower visitation rate and offspring weight decreased as the magnitude of asynchrony increased. Due to low offspring production and a lack of female offspring across treatments, we were unable to assess the effects of phenological asynchrony on total offspring produced or sex ratio. Findings suggest that post-emergence food deprivation caused by phenological asynchrony may affect offspring size by influencing nest-provisioning rates. In solitary bees, body size influences wintering survival, fecundity, and mating success. Thus, phenological asynchrony may have consequences for solitary bee populations that stem from reduced flower visitation rates, and these consequences may increase as the magnitude of asynchrony increases. Because many wild flowering plants and crops rely on pollination services provided by bees for reproductive success, bee responses to phenological asynchrony may also affect wild plant biodiversity and crop yields.     

Nurse bees support the physiological development of young bees (Apis mellifera L.)

Newly emerged worker honeybees (focal bees) were caged individually for 8 days either isolated or together with one companion bee of known age (2–30 days) taken from a colony. The companion bee was replaced every 2nd day. After 8 days, various parameters were investigated in the focal bees as indicators of the level of development. Focal bees which had been caged with 6-day-old companion bees were better developed than isolated focal bees, newly emerged bees, or focal bees caged with almost all other ages of companion bees. They had hypopharyngeal glands that were larger and contained more protein, their thoraces had a higher protein content, and they had a higher rate of proteolytic activity in the midgut. Although the focal bees were supplied with pollen as well as honey, they consumed only small amounts of pollen. We attribute their better development to their having been fed worker jelly by the accompanying companion bees. The 6-day-old companion bees consumed high quantities of pollen and spent more time (18.7 ± 11.85 s/h) feeding focal bees than 12-day-old bees (6.5 ± 4.09 s/h) or foragers (no feeding of focal bees). The results show that even under such artificial conditions, the exchange of food (trophallaxis) promotes the development of young honeybee workers. Accepted: 26 February 1999     

Influence of Pollen Nutrition on Honey Bee Health: Do Pollen Quality and Diversity Matter?

Honey bee colonies are highly dependent upon the availability of floral resources from which they get the nutrients (notably pollen) necessary to their development and survival. However, foraging areas are currently affected by the intensification of agriculture and landscape alteration. Bees are therefore confronted to disparities in time and space of floral resource abundance, type and diversity, which might provide inadequate nutrition and endanger colonies. The beneficial influence of pollen availability on bee health is well-established but whether quality and diversity of pollen diets can modify bee health remains largely unknown. We therefore tested the influence of pollen diet quality (different monofloral pollens) and diversity (polyfloral pollen diet) on the physiology of young nurse bees, which have a distinct nutritional physiology (e.g. hypopharyngeal gland development and vitellogenin level), and on the tolerance to the microsporidian parasite Nosema ceranae by measuring bee survival and the activity of different enzymes potentially involved in bee health and defense response (glutathione-S-transferase (detoxification), phenoloxidase (immunity) and alkaline phosphatase (metabolism)). We found that both nurse bee physiology and the tolerance to the parasite were affected by pollen quality. Pollen diet diversity had no effect on the nurse bee physiology and the survival of healthy bees. However, when parasitized, bees fed with the polyfloral blend lived longer than bees fed with monofloral pollens, excepted for the protein-richest monofloral pollen. Furthermore, the survival was positively correlated to alkaline phosphatase activity in healthy bees and to phenoloxydase activities in infected bees. Our results support the idea that both the quality and diversity (in a specific context) of pollen can shape bee physiology and might help to better understand the influence of agriculture and land-use intensification on bee nutrition and health.     

Molecular assays of pollen use consistently reflect pollinator visitation patterns in a system of flowering plants

Determining how pollinators visit plants vs. how they carry and transfer pollen is an ongoing project in pollination ecology. The current tools for identifying the pollens that bees carry have different strengths and weaknesses when used for ecological inference. In this study we use three methods to better understand a system of congeneric, coflowering plants in the genus Clarkia and their bee pollinators: observations of plant–pollinator contact in the field, and two different molecular methods to estimate the relative abundance of each Clarkia pollen in samples collected from pollinators. We use these methods to investigate if observations of plant–pollinator contact in the field correspond to the pollen bees carry; if individual bees carry Clarkia pollens in predictable ways, based on previous knowledge of their foraging behaviors; and how the three approaches differ for understanding plant–pollinator interactions. We find that observations of plant–pollinator contact are generally predictive of the pollens that bees carry while foraging, and network topologies using the three different methods are statistically indistinguishable from each other. Results from molecular pollen analysis also show that while bees can carry multiple species of Clarkia at the same time, they often carry one species of pollen. Our work contributes to the growing body of literature aimed at resolving how pollinators use floral resources. We suggest our novel relative amplicon quantification method as another tool in the developing molecular ecology and pollination biology toolbox.     

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.

     

Observations on the initiation and stimulation of oviposition of theVarroa mite

FemaleVarroa mites were collected from adult bees and were classified as swollen or not swollen. After introduction of these mites into recently sealed worker brood cells the average number of offspring of reproducing swollen mites was similar to that of naturally invaded mites, but the non-swollen mites produced a significantly lower number of offspring. This suggests that the oviposition of adult mites is stimulated by a preceding stay on adult bees. When (non-swollen) mites collected from brood cells were kept for 35 days in Eppendorf test tubes containing a larva or a pupa, their reproduction was similar to that of swollen mites.Contact of young mites, collected from brood cells, with adult bees was not essential for the initiation of oviposition. However, the number of offspring of reproducing mites, even after a third or fourth introduction into brood cells, was as a rule lower than that of mites that had been in contact with adult bees.The period of artificial introduction into sealed brood cells proved to be essential for subsequent reproduction. When introduced 48–52 h or 72–76 h after cell sealing, the mites did not produce eggs. When introduced 0–4 h after cell sealing a high percentage of the mites reproduced. Contact of the mites with a spinning larva seems necessary for initiation of oviposition.     

Pollen consumed by the solitary bee Tetrapedia diversipes (Apidae: Tetrapediini) in a tropical agroecosystem

Pollen analysis of the larval food supply is an important tool for identifying the plants that provide the floral resources used by bees. The present study documents the pollen sources consumed by larvae of the solitary bee Tetrapedia diversipes in a tropical agroecosystem. A total of 60 pollen types were recorded with three families being the most important. Euphorbiaceae (60.5%), Malpighiaceae (16.8%) and Asteraceae (12.2%) pollen had the greatest representation in the samples examined. The pollen of Dalechampia dioscoreifolia predominated in the diet of the larvae of T. diversipes (RF?=?56.35%) and indicates the importance of this plant in maintaining populations of this solitary bee.     

Notes on the pollination mechanisms ofMoraea inclinata andM. brevistyla (Iridaceae)

Individual flowers ofMoraea inclinata are nectariferous and last about six hours. They appear to be pollinated largely by bees in the familyHalictidae (Lasioglossum spp.,Nomia spp.,Zonalictus) and to a lesser extent by bees in the familyAnthophoridae (Amegilla). The mechanism of bee-pollination inM. inclinata is the Iris type; i.e., each flower consists of three pollination units (an outer tepal, a partly exserted anther, and the opposed style branch which terminates in a pair of petal-like crests). Bees rarely visit more than one pollination unit per flower. Transferral of pollen to the bee is passive and nototribic although all bees collected on the flowers were female and 55% of the bees carried pollen loads with 2–5 pollen taxa in their scopae.Moraea brevistyla flowers are nectariferous but lack scent and last two days. They are visited infrequently by bees and only one femaleLasioglossum spec. carried the pollen ofM. brevistyla. Unlike flowers ofM. inclinata those ofM. brevistyla deposit pollen only on the head and thorax. Bee-mediated autogamy in both species is avoided due to the erratic foraging patterns of the bees and the flexibility of each stigma lobe as the bee backs out of the flower. Approximately 2–4 flowers in the inflorescences of both species (6–8 flowers/infloresence) develop into capsules.     

Cryptic species and hidden ecological interactions of halictine bees along an elevational gradient

Changes of abiotic and biotic conditions along elevational gradients represent serious challenges to organisms which may promote the turnover of species, traits and biotic interaction partners. Here, we used molecular methods to study cuticular hydrocarbon (CHC) profiles, biotic interactions and phylogenetic relationships of halictid bees of the genus Lasioglossum along a 2,900 m elevational gradient at Mt. Kilimanjaro, Tanzania. We detected a strong species turnover of morphologically indistinguishable taxa with phylogenetically clustered cryptic species at high elevations, changes in CHC profiles, pollen resource diversity, and a turnover in the gut and body surface microbiome of bees. At high elevations, increased proportions of saturated compounds in CHC profiles indicate physiological adaptations to prevent desiccation. More specialized diets with higher proportions of low‐quality Asteraceae pollen imply constraints in the availability of food resources. Interactive effects of climatic conditions on gut and surface microbiomes, CHC profiles, and pollen diet suggest complex feedbacks among abiotic conditions, ecological interactions, physiological adaptations, and phylogenetic constraints as drivers of halictid bee communities at Mt. Kilimanjaro.     

Suitability of pollen sources for the development and reproduction of Coleomegilla maculata (Coleoptera: Coccinellidae) under simulated drought conditions

Laboratory experiments compared the nutritive value of various pollen sources for the development of Coleomegilla maculata DeGeer under conditions of continuous water availability and simulated drought. When water was continuously available, larval survival was not different from 100% on diets of frozen eggs of Ephestia kuehniella Zeller, corn pollen, sorghum pollen, or pulverized bee pollen, whereas survival of larvae was significantly reduced on the latter three diets in the simulated drought treatment. Pollen of cultivated sunflower, Helianthus annus L., proved fatal to both larvae and adults; its surface structure caused clumping and accumulation on the insect cuticle that led to death from exhaustion/desiccation in petri dishes. The Ephestia egg diet yielded shorter developmental times and heavier adult weights than any pollen diet in both treatments. The drought treatment increased developmental time on all diets with a significant treatment–diet interaction. Drought reduced the adult weight of females on the sorghum pollen diet, and that of both sexes on the bee pollen diet, again with a significant treatment–diet interaction. Initial water content was highest in corn pollen (36.8%), followed by Ephestia eggs (29.2%), sorghum pollen (25.3%), sunflower pollen (8.7%), and bee pollen (4.6%), but did not appear correlated with C. maculata larval survival on pollen sources under drought conditions. Reproductive adult females that received corn or sorghum pollen as a supplement to Ephestia eggs did not differ in fecundity or fertility from those fed only Ephestia eggs.     

Influence of pollen origin on performance of Bombus terrestris micro-colonies

The effect of the floral origin of pollen on the reproductive success of Bombus terrestris Latrum bumblebee (Apidae: Bombinae), was investigated by feeding micro-colonies of queenless workers with different pollen types. We used a commercial pollen blend and three unifloral pollens, Prunus, Salix, and Taraxacum. Among the unifloral pollens, pollen quality did not influence egg production, but did influence egg laying delay and larval growth. The quality of pollens varied according to their protein content and protein efficacy (PE). Pollen from Prunus (27.5% w/w protein, PE=10) resulted in the largest number of progeny, whereas Taraxacum (17.2% w/w protein, PE=0) did not result in any offspring, due to high oophagy and larval ejection. Salix (20% w/w protein) and the blend (22.8% protein) diets gave rise to intermediate reproductive outputs. When pollen quality was sufficient for larval growth, the fitness of the male offspring was not affected over the range of the experimental diets. Our results suggest that quantitative and qualitative variations of pollen proteins have considerable influence on the reproductive success of bumblebees. Furthermore, larval growth has specific nutritive demands not provided by Taraxacum pollen, which is missing two essential amino acids.     

The interaction between pollinator size and the bristle staminode of Penstemon digitalis (Scrophulariaceae)

Penstemon digitalis, a prairie species whose flowers possess a large bristle staminode, is visited by eight bee species of varying size. Two sets of field experiments involving staminode removal were performed to test pollination efficiency in relation to bee size. Our data indicate that bristle staminode presence and function are influenced by size-dependent selection on bee body size and associated pollen transporting attributes. The first experiment compared staminode presence and removal in open-pollinated flowers. Staminode removal significantly reduced total pollen deposition but had no effect on total pollen removal. The second experiment utilized single bee visits to assess the interaction between pollinator size and staminode presence on the rate of pollen deposition and removal. This experiment indicated that staminode removal resulted in fewer pollen grains deposited on stigmas and less pollen removed from anthers for both large and small bees (the contrary was true for medium bees). Although the number of pollen grains deposited was greatly reduced for large bees, staminode removal reduced deposition efficiency most notably for small bees by 41.3%, reducing female reproductive success. Staminode removal increased pollen removal efficiency most notably for medium bees by 22.7%, reducing male reproductive success. Mechanisms of staminode function are discussed.     

Effect of protein source in diets fed during gestation and lactation on food intake regulation in male offspring of Wistar rats

We hypothesized that protein source in the nutritionally adequate AIN-93G diets fed during gestation, lactation, and weaning influences food intake (FI) regulation in male offspring of Wistar rats. Pregnant rats were fed the recommended casein-based (C) or soy protein-based (S) diet during gestation (experiment 1) or during gestation and lactation (experiment 2). Pups (n = 12 per group) weaned to C or S diets were followed for 9 wk (experiment 1) or 14 wk (experiment 2). At termination, body weight was 5.4% and 9.4% higher, respectively, in offspring of dams fed the S diet. Altered FI regulation was shown by failure of devazepide (a CCK-A receptor blocker) to block FI reduction after protein preloads in offspring of S diet-fed dams, whereas it had a strong effect on offspring of C diet-fed dams (P < 0.005). Similarly, naloxone (an opioid receptor blocker) blocked FI reduction more after casein than after soy protein preloads (P < 0.01). In experiment 2, offspring of dams fed the S diet had higher hypothalamic gene expression of agouti related protein at weaning (P < 0.05), and higher FI was found throughout postweaning (P < 0.0001). FI reduction after protein preloads at week 7 and after glucose preloads at week 13 was greater in offspring of C diet-fed dams (P < 0.05). Plasma insulin at weaning and insulin, ghrelin, and glucagon-like peptide-1 at week 15 were higher in offspring of S diet-fed dams (all P < 0.05). In conclusion, nutritionally complete C and S diets consumed during gestation and lactation differ in their effects on body weight and FI regulation in the offspring. Extending the diet from gestation alone to throughout gestation and lactation exaggerated the adverse effects of the S diet. However, the diet consumed postweaning had little effect on the outcome.     

Diet characterisation of solitary bees on farmland: dietary specialisation predicts rarity

Changes in agricultural practice across Europe and North America have been associated with declines in wild bee populations. Bee diet breadth has been associated with sensitivity to agricultural intensification, but much of this analysis has been conducted at the categorical level of generalist or specialist, and it is not clear to what extent the level of generalisation within generalist species is also associated with species persistence. We used pollen load analysis to quantify the pollen diets of wild solitary bees on 19 farms across southern England, UK. A total of 72 species of solitary bees were recorded, but only 31 species were abundant enough to allow for formal diet characterisation. The results broadly conformed to existing literature with the majority of species polylectic and collecting pollen from a wide range of plants. Pollen load analysis consistently identified pollens from more plant species and families from each bee species than direct observation of their foraging behaviour. After rarefaction to standardise pollen load sample sizes, diet breadth was significantly associated with frequency of occurrence, with more generalist bees present on more farms than less generalist bees. Our results show that the majority of bee species present on farmland in reasonable numbers are widely variable in their pollen choices, but that those with the broadest diet were present on the greatest number of farms. Increasing the diversity of plants included in agri-environment schemes may be necessary to provide a wider range of pollen resources in order to support a diverse bee community on farmland.     

Morphological specialization of heterantherous Rhynchanthera grandiflora (Melastomataceae) accommodates pollinator diversity

• The tropical Melastomataceae are characterized by poricidal anthers which constitute a floral filter selecting for buzz‐pollinating bees. Stamens are often dimorphic, sometimes with discernible feeding and pollinating functions. Rhynchanthera grandiflora produces nectarless flowers with four short stamens and one long stamen; all anthers feature a narrow elongation with an upwards facing pore.
• We tested pollen transfer by diverse foraging bees and viability of pollen from both stamen types. The impact of anther morphology on pollen release direction and scattering angle was studied to determine the plant's reproductive strategy.
• Medium‐sized to large bees sonicated flowers in a specific position, and the probability of pollen transfer correlated with bee size even among these legitimate visitors. Small bees acted as pollen thieves or robbers. Anther rostrum and pore morphology serve to direct and focus the pollen jet released by floral sonication towards the pollinator's body. Resulting from the ventral and dorsal positioning of the short and long stamens, respectively, the pollinator's body was widely covered with pollen. This improves the plant's chances of outcrossing, irrespective of which bee body part contacts the stigma. Consequently, R. grandiflora is also able to employ bee species of various sizes as pollen vectors.
• The strategy of spreading pollen all over the pollinator's body is rather cost‐intensive but counterbalanced by ensuring that most of the released pollen is in fact transferred to the bee. Thus, flowers of R. grandiflora illustrate how specialized morphology may serve to improve pollination by a functional group of pollinators.