Nectar usage in a southern Arizona hawkmoth community

Abstract.  1. Hawkmoths (Sphingidae) are important plant associates at two life-history stages: larvae are herbivorous, whereas adults are nectar feeders and often pollinators. The diversity and identities of plants used for nectar is poorly known, however.
2. This study takes a community-level approach to hawkmoth nectar plant usage in a semi-arid grassland habitat in southern Arizona, U.S.A.
3. Pollen carried on the proboscis was identified from over 700 individuals of 14 hawkmoth species attracted to lights over a 2-year period.
4. Two plant species dominated pollen loads, suggesting that hawkmoths use these species extensively as nectar sources: Datura wrightii (Solanaceae), a classic hawkmoth-pollinated plant, and Agave palmeri (Agavaceae), which is known to be used extensively by bats. Field surveys indicate that both species are relatively rare in the flowering community. Little or no pollen was present on the moths from the most common plant species in flower during the study.
5. The dominance of Agave in pollen loads suggests that this typically bat-pollinated species may be subsidising pollinator populations of the hawkmoth-pollinated flora.
6. Three groups of hawkmoths within this community are identified based on larval diets (reported in the literature) and adult diets (documented here): those that, at a given site, heavily exploit the same plant species at both life-history stages ( Manduca sexta and M. quinquemaculata ); those that have broad local associations at both life-history stages ( Hyles lineata ); and those that exhibit narrow but non-overlapping local associations at the two life-history stages (all other hawkmoths at this site).     

Nectarivory by the plant-tissue feeding predator Macrolophus pygmaeus Rambur (Heteroptera: Miridae): Nutritional redundancy or nutritional benefit?

Most predators and parasitoids feed on plant-provided food (nectar, pollen) or engage in herbivory during at least part of their life stages. Plant feeding by these insects plays an important role in driving predator-herbivore dynamics. Thus, understanding the effects of plant feeding on omnivores could be an important element in improving biological control strategies. The mirid Macrolophus pygmaeus is an omnivorous heteropteran predator of whitefly and other pests. Unlike other predators that need to seek out accessible nectar to meet their carbohydrate requirements, mirid bugs can access the plant's carbohydrate resources by feeding directly on plant tissues. Leaf and stem feeding could be seen as a nutritional surrogate that allows mirids to become independent of nectar availability. However, to date feeding experiments have not yet considered nectar feeding by these mirid predators. In this study we demonstrate that M. pygmaeus survival is prolonged on broad bean plants featuring extrafloral nectar as compared to broad bean with extrafloral nectaries removed, irrespective of the presence of cattail pollen. Survival on extrafloral nectar was comparable to the survival by individuals kept on broad bean provided with eggs of Ephestia kuehniella as prey. Also, a greater proportion of mirid females laid eggs when extrafloral nectar was available as compared to those confined on nectariless plants without supplemental food.     

Studies in Garcinia, dioecious tropical forest trees: the phenology, pollination biology and fertilization of G. hombroniana Pierre

RICHARDS, A. J., 1990. Studies in Garcinia , dioecious tropical forest trees: the phenology, pollination biology and fertilization of G. hombroniana Pierre . Garcinia hombroniana is a facultative agamosperm which is pollinated by Trigona bees. Nectar is restricted to the large discoid stigma (or pistillode in male flowers), which also captures and hydrates pollen. The 'wet' stigma and binucleate pollen suggest that Garcinia arose from hermaphrodite plants with a gametophytic self-incompatibility system.
On stigmas, nectar is secreted early on three or four successive days. On male pistillodes, nectar is secreted when anthers dehisce, on the second morning after anthesis. Pollen is most viable when freshly collected, but some viability remains four days after collection. Pollen germinates within 24 h of hydration. Similar results to pollinations are obtained by germinating pollen in 1 % sucrose.
Garcinia hombroniana flowers principally from January to June. Cultivated females are considered as 'big bang' strategists. Male flowers are considered as 'steady state' strategists.     

Bee visits and the nectar of Echium vulgare L. and Sinapis alba L.

Abstract. 1. The time-course of anthesis of Echium vulgare is described.
2. Diel changes in the sugar concentration of the nectar, the quantity of nectar and the quantity of sugar per flower are illustrated for E.vulgare and for Sinapis alba.
3. These changes are interpreted in terms of (a) the periodicity of secretion and (b) the influence of microclimate and insect visits on post-secretory changes in the composition and volume of nectar.
4. There was hour-to-hour and day-to-day variation in the species composition and the proportion of workers taking nectar rather than nectar plus pollen among the social bees visiting E.vulgare.
5. Honeybee visits to E.vulgare were more numerous in humid weather, when there was enough nectar per flower for their relatively short tongues to reach, and in an area where the corollas grew shorter than they did elsewhere.
6. The significance of changes in the caloric content, volume, concentration, viscosity and sugar composition is discussed from the points of view of insects and ecologists.     

Correlation between octopaminergic signalling and foraging task specialisation in honeybees

Regulation of pollen and nectar foraging in honeybees is linked to differences in the sensitivity to the reward. Octopamine (OA) participates in the processing of reward-related information in the bee brain, being a candidate to mediate and modulate the division of labour among pollen and nectar foragers. Here we tested the hypothesis that OA affects the resource preferences of foragers. We first investigated whether oral administration of OA is involved in the transition from nectar to pollen foraging. We quantified the percentage of OA-treated bees that switched from a sucrose solution to a pollen feeder when the sugar concentration was decreased experimentally. We also evaluated if feeding the colonies sucrose solution containing OA increases the rate of bees collecting pollen. Finally, we quantified OA and tyramine (TYR) receptor genes expression of pollen and nectar foragers in different parts of the brain, as a putative mechanism that affects the decision-making process regarding the resource type collected. Adding OA in the food modified the probability that foragers switch from nectar to pollen collection. The proportion of pollen foragers also increased after feeding colonies with OA-containing food. Furthermore, the expression level of the AmoctαR1 was upregulated in foragers arriving at pollen sources compared with those arriving at sugar-water feeders. Using age-matched pollen and nectar foragers that returned to the hive, we detected an upregulated expression of a TYR receptor gene in the suboesophageal ganglia. These findings support our prediction that OA signalling affects the decision in honeybee foragers to collect pollen or nectar.     

Pollen amino acids—an additional diet for a nectar feeding butterfly?

The increase of the amino acid concentration over different time intervals in artificial nectar (i.e., a sucrose solution) due to pollen contamination was investigated in four Californian plant species (Aesculus californica, Amsinckia lunaris, Brodiaea pulchella, Carduus pycnocephalus), which are important nectar resources for a Californian colony of the butterflyBattus philenor as well as for other insects. The increase of the amino acid concentration in the medium is different in all four species and seems to be determined by a variety of factors including permeability of the pollen grain wall and presence or absence of pores. The results suggest a passive diffusion process of the free pollen amino acids into the medium rather than an active release. Implications from the experiments forBattus philenor and for other nectar feeding pollinators are discussed. A possible complementary effect of free pollen and nectar amino acids is proposed for plant species in which pollen is likely to be knocked into nectar by their flower visitors. A possible evolutionary pathway from nectar feeding butterflies such asBattus philenor to the complex derived pollen feeding habit in theHeliconius butterflies is proposed.     

On the function of pea flower feeding by Bruchus pisorum

Pea flower feeding by adult pea weevils, Bruchus pisorum (L.) (Coleoptera: Bruchidae), with special emphasis on nectar feeding, was investigated in a series of laboratory experiments. Male and female adults robbed nectar from flowers of the garden and field pea, Pisum sativum L., and females which fed on the nectar, petals, and female organs of pea flowers lived significantly longer than those denied food and water and those that fed on water only. The results of other experiments suggested that pea flower qualities other than pollen influenced the reproductive success of female B. pisorum. It is hypothesized that pollen seeking B. pisorum effected cross-pollination in the wild progenitor of the modern-day autogamous pea, and adult pea weevils of both sexes rob pea nectar to obtain a readily available source of energy to sustain flight.     

Evolution of the suctorial proboscis in pollen wasps (Masarinae, Vespidae)

The morphology and functional anatomy of the mouthparts of pollen wasps (Masarinae, Hymenoptera) are examined by dissection, light microscopy and scanning electron microscopy, supplemented by field observations of flower visiting behavior. This paper focuses on the evolution of the long suctorial proboscis in pollen wasps, which is formed by the glossa, in context with nectar feeding from narrow and deep corolla of flowers. Morphological innovations are described for flower visiting insects, in particular for Masarinae, that are crucial for the production of a long proboscis such as the formation of a closed, air-tight food tube, specializations in the apical intake region, modification of the basal articulation of the glossa, and novel means of retraction, extension and storage of the elongated parts. A cladistic analysis provides a framework to reconstruct the general pathways of proboscis evolution in pollen wasps. The elongation of the proboscis in context with nectar and pollen feeding is discussed for aculeate Hymenoptera.     

Size-related selection of food plants by bumblebees

Abstract. 1. A positive correlation between the tongue length of conspecific workers collecting nectar from seven plant species and the corolla length of the flowers probed was found for B.lapidarius and B.pascuomm but not B.terrestris . No simple relationship was found between the volume, sugar weight or concentration of nectar in flowers and the tongue or wing length of probing bees.
2. B.terrestris workers collecting pollen from four plant species producing pollen only, were found to differ in size according to the type of pollen presentation mechanism and the pollen content per flower. Body size variation was also related to the foraging of pollen plus nectar from two other plant species.     

Mid-Cretaceous charred fossil flowers reveal direct observation of arthropod feeding strategies

Although plant-arthropod relationships underpin the dramatic rise in diversity and ecological dominance of flowering plants and their associated arthropods, direct observations of such interactions in the fossil record are rare, as these ephemeral moments are difficult to preserve. Three-dimensionally preserved charred remains of Chloranthistemon flowers from the Late Albian to Early Cenomanian of Germany preserve scales of mosquitoes and an oribatid mite with mouthparts inserted into the pollen sac. Mosquitoes, which today are frequent nectar feeders, and the mite were feeding on pollen at the time wildfire consumed the flowers. These findings document directly arthropod feeding strategies and their role in decomposition.     

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.     

Influence of floral resources on sugar feeding and nutrient dynamics of a parasitoid in the field

Abstract.  1. To investigate how floral resources impact feeding and nutrient dynamics of parasitoids, the ichneumonid Diadegma insulare (Cresson) was monitored in cabbage fields with and without buckwheat Fagopyrum esculentum Moench borders. Biochemical analyses of field-collected wasps determined whether sugar feeding occurred, and whether it enhanced nutrient levels and increased parasitism of diamondback larvae Plutella xylostella (L.) from 2000 to 2003. The effects of experimental spatial scale on sugar feeding dynamics were also evaluated in 2001.
2. The majority of D. insulare foraging on cabbage had fed upon sugar, but floral borders did not consistently increase the proportion of sugar-fed wasps or their levels of glycogen and lipid over all 4 years. Floral borders increased sugar storage by females in 2001. More females fed on sugar when experimental plots were separated by at least 800 m compared with 67 m.
3. Feeding on buckwheat nectar vs. honeydew produced by the soybean aphid Aphis glycines Matsumura in surrounding fields was distinguished using the ratio of fructose to total sugars, which was higher in nectar- than in honeydew-fed parasitoids. Floral borders increased the probability that females collected from adjacent crops were nectar-fed in 2002, demonstrating that floral resources were utilised by foraging parasitoids. Diadegma insulare that fed upon any sugar source had higher levels of sugar and glycogen than unfed wasps, and feeding on honeydew appeared to enhance nutrient storage.
4. The prevalence of sugar or nectar feeding by female D. insulare was not correlated with parasitism rates over the 4 years from all plots, but nectar feeding was positively correlated with parasitism rates within floral plots.     

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.     

Energy regulation by traplining hummingbirds

1. A published model of constant diurnal energy accumulation by territorial hummingbirds does not accurately reflect the temporal distribution of feeding behaviour of traplining hummingbirds, Phaethornis longirostris (Long-Tailed Hermit Hummingbirds).
2. In an enclosure study, gross nectar intake by P . longirostris decreased through the day, mirroring nectar production rates in its natural food-flowers and mimicking its natural foraging patterns.
3. Using a simulation model, the energetic consequences of constant and decreasing net energy intake rates for traplining hummingbirds are compared.
4. Given natural patterns of nectar production, model birds with decreasing diurnal net intake rates met their energetic needs with fewer flowers than those with constant net intake, and spent less time foraging.
5. It is concluded that P . longirostris do not satisfy the physiological assumptions of the published model, and that in this way they are different from the territorial species on which the model has previously been tested.     

Adult pollen diet essential for egg maturation by a solitary Osmia bee

Reproduction is a nutritionally costly activity for many insects, as their eggs are rich in lipids and proteins. That cost seems especially acute for non-social bees, which for their size, lay enormous eggs. All adult female bees visit flowers, most of them to collect pollen and nectar, or sometimes oils, to feed their progeny. For adult bees, the need for pollen feeding has only been detailed for the honey bee, Apis mellifera. To experimentally test for the reproductive value of adult pollen feeding by a non-social bee, Osmia californica (Hymenoptera: Apiformes: Megachilidae), young female bees plus males were released into large glasshouse cages provided with either a male-fertile sunflower cultivar or a pollen-less one. Females regularly visited and drank nectar from flowers of both cultivars. Abundant orange pollen was seen regularly in guts of females confined with the male-fertile sunflowers, indicative of active pollen ingestion. All females’ terminal oocytes (next egg to be laid) were small at emergence. Oocytes of females confined with the pollen-less sunflowers remained small, despite frequent nectaring and exposure to other floral stimuli. In contrast, the basal oocytes of female O. californica with access to pollen had swelled to full size within ten days following emergence, enabling them to lay eggs in provided nest tubes. Adult females of this solitary bee required dietary pollen to reproduce; nitrogen stores acquired as larvae were inadequate. Early and regular pollen feeding in part paces the onset and maximum tempo of solitary bees’ lifetime reproductive output.     

Economic motivation for plant species preferences of pollen-collecting bumble bees

1. Pollen is an essential resource for bees as it provides the proteins needed for larval development and growth. Bumble bees typically collect pollen from a limited subset of the available plant species, indicating that bees perceive qualitative and/or quantitative differences between species. Because bumble bees typically exhibit different foraging behaviours when collecting pollen than when collecting nectar, and often visit different plant species, the economics of pollen collection are subject to unique benefits and costs.
2. The influences of pollen standing crop, grain volume, protein content and foraging costs on the choice of plant species by pollen-collecting bumble bees were assessed. Pollen was collected from different plant species available to bees and these species were ranked according to eleven currencies that differentially incorporate these influences on foraging economics.
3. Comparison of observed preferences at three sites with species rankings based on foraging currencies revealed that bumble bees do not assess plant species based solely on intrafloral characteristics. Instead, pollen-foraging bumble bees are sensitive to protein availability at the site as a whole as well as foraging costs, and forage in a manner that maximizes the site-specific efficiency of protein collection (protein collected/energetic costs). Such behaviour would promote proper larval development while maximizing forager lifespan.     

How much pollen can thrips destroy?

ABSTRACT. 1. A laboratory technique for measuring the number of pollen grains consumed by thrips is described.
2. Thrips imaginis Bagnall and Thrips obscuratus (Crawford) (Thripidae) were studied particularly on pollen of the kiwifruit Actinidia deliciosa (A. Chevalier) Liang & Ferguson in New Zealand, and Echium plantagineum L. in Australia.
3. Mean daily feeding rates (in grains per thrips per day) ranged from 29 to 843, with an individual rate as high as 1626 for T.imaginis larvae II on E. plantagineum.
4. The time taken to feed on a single grain was proportional to grain volume, and decreased with temperature.
5. Daily feeding rates were significantly different between pollens, and were higher for smaller grains. The total volume of pollen contents consumed and the total time spent ingesting this volume per thrips per day may be constant with respect to pollen species.
6. Daily feeding rates were equivalent to 0.2–0.7% of the average total pollen production of a flower per thrips per day.
7. Extrapolation of the daily feeding rates suggests that pollen damage by thrips could sometimes be reducing crop yield or plant fitness.     

Pollen feeding proteomics: Salivary proteins of the passion flower butterfly,Heliconius melpomene

While most adult Lepidoptera use flower nectar as their primary food source, butterflies in the genus Heliconius have evolved the novel ability to acquire amino acids from consuming pollen. Heliconius butterflies collect pollen on their proboscis, moisten the pollen with saliva, and use a combination of mechanical disruption and chemical degradation to release free amino acids that are subsequently re-ingested in the saliva. Little is known about the molecular mechanisms of this complex pollen feeding adaptation. Here we report an initial shotgun proteomic analysis of saliva from Heliconius melpomene. Results from liquid-chromatography tandem mass-spectrometry confidently identified 31 salivary proteins, most of which contained predicted signal peptides, consistent with extracellular secretion. Further bioinformatic annotation of these salivary proteins indicated the presence of four distinct functional classes: proteolysis (10 proteins), carbohydrate hydrolysis (5), immunity (6), and “housekeeping” (4). Additionally, six proteins could not be functionally annotated beyond containing a predicted signal sequence. The presence of several salivary proteases is consistent with previous demonstrations that Heliconius saliva has proteolytic capacity. It is likely that these proteins play a key role in generating free amino acids during pollen digestion. The identification of proteins functioning in carbohydrate hydrolysis is consistent with Heliconius butterflies consuming nectar, like other lepidopterans, as well as pollen. Immune-related proteins in saliva are also expected, given that ingestion of pathogens is a likely route to infection. The few “housekeeping” proteins are likely not true salivary proteins and reflect a modest level of contamination that occurred during saliva collection. Among the unannotated proteins were two sets of paralogs, each seemingly the result of a relatively recent tandem duplication. These results offer a first glimpse into the molecular foundation of Heliconius pollen feeding and provide a substantial advance towards comprehensively understanding this striking evolutionary novelty.     

Forage quality and quantity affect red mason bees and honeybees differently in flowers of strawberry varieties

Nectar is a vital source of energy for bees and other pollinators and pollen represents the only source of protein in the diet of bees. Nectar and pollen quality and quantity can therefore affect foraging choices. Strawberry, Fragaria × ananassa (Rosaceae), is a flowering crop that requires insect pollination for the berries to develop optimally. The solitary red mason bee, Osmia bicornis L. (Hymenoptera: Megachilidae), occurs naturally but like the eusocial western honeybee, Apis mellifera mellifera L. (Hymenoptera: Apidae), it is also a commercially reared pollinator used in strawberry production. We hypothesized that strawberry nectar and pollen quality would affect the foraging choice of these two types of bees. In this study nectar and pollen quality is represented by various levels of sugar and protein content, respectively, as well as the number of open strawberry flowers in the experimental field, would affect the foraging choice of these two types of bees. Consistent with previous studies, we found significant and major differences between strawberry varieties in proportions of sucrose in the nectar sugar and in pollen viability – a proxy for pollen protein content. All measured parameters had a significant effect on red mason bee visitation frequency. Contrary to expectations, honeybee foraging behavior was only affected by the number of open flowers and not by any of the quality parameters measured. Our findings indicate that red mason bees were capable of assessing nectar and pollen quality and prioritize accordingly. The pattern observed indicates that individual red mason bees changed foraging focus between strawberry varieties depending on whether nectar or pollen was collected. Our results suggest that targeted breeding of varieties toward high levels of nectar sugar and sucrose concentrations and high pollen protein content may increase pollination success from red mason bees and possibly other solitary bees.     

Field energetics and the estimation of pollen and nectar intake in the marsupial honey possum, Tarsipes rostratus, in heathland habitats of South-Western Australia

A method is described, based on the simultaneous turnover of both stable (¹⁸O) and radioactive isotopes (³H and ²²Na), whereby the daily nectar and pollen intake of free-ranging marsupial honey possums (Tarsipes rostratus) may be estimated. The field metabolic rate is measured using doubly labelled water and nectar intake is estimated independently from the measured water and sodium fluxes. The method assumes that free-water intake is negligible (but may be accounted for if not the case), that virtually all dietary sodium is derived from nectar rather than from pollen, and that the animals are in energetic balance over the period of measurement. These assumptions have been tested and found to be robust, except during periods of heavy rain when significant intakes of free-water were recorded. Leaching experiments with pollen grains suggest that less than 10% of the sodium ingested by honey possums is derived from pollen and calculations thus assumed a 90%:10% split between nectar and pollen. Nectar intake averaged 5.9 ± 0.6 ml · day⁻¹ and regressing nectar intake on daily change in body mass predicts an intake of approximately 7 ml · day⁻¹ nectar to maintain balance for a 9 g honey possum. Estimates of pollen intake averaged 660 ± 156 mg · day⁻¹ and a similar regression analysis of the data predicts that a daily intake of approximately 1 g pollen would be needed to maintain mass balance of honey possums. Estimated nectar and pollen intakes did not differ significantly between males and females, but nectar intake was higher in winter compared with dry periods of the year. The sugar content of nectar falls during winter, however, and the overall energy derived from nectar thus remains roughly constant. Estimates of pollen and nectar intake for individual animals were not significantly correlated, suggesting that honey possums forage selectively for these two food items. Accepted: 19 August 1999