Feeding responses of adult butterflies, Nymphalis xanthomelas, Kaniska canace and Vanessa indica, to components in tree sap and rotting fruits: synergistic effects of ethanol and acetic acid on sugar responsiveness

Exuded tree sap and rotting fruits are important feeding sources for adults of many butterfly species. However, the chemical composition of such rotting foods and the effects of these constituents on butterfly feeding behaviour have rarely been investigated. Chemical analyses revealed that these food sources contain several major components, including fructose, glucose, ethanol and acetic acid. Moreover, 15 amino acids were identified from the exuded tree sap. The gustatory responsiveness to these components was examined in the proboscis of nymphalid adults of three species (Nymphalis xanthomelas, Kaniska canace and Vanessa indica). The first two species primarily feed on sap and rotting fruits, while V. indica frequently visits flowers. Fructose elicited larger feeding responses from these saprophagous butterflies than glucose, which had a feeding-stimulatory effect only on N. xanthomelas at a concentration equivalent to that of the food. Ethanol, acetic acid and the five major amino acids identified in tree sap did not elicit feeding responses by themselves. However, ethanol and/or acetic acid, when mixed with these sugars at equivalent concentrations, evoked feeding and enhanced probing responses in K. canace and V. indica. These results suggest that ethanol and acetic acid, together with sugars at low concentrations, synergistically stimulate butterfly feeding behaviour.     

Divergent behavioral and electrophysiological taste responses in the mid-legs of adult butterflies, Vanessa indica and Argyreus hyperbius

Adult nymphalid butterflies possess sensilla trichodea (ST) that perceive taste in their walking legs. We examined whether the gustatory responses to mid-leg tarsal stimulation were different between Vanessa indica (rotting-food feeder) and Argyreus hyperbius (flower-nectar feeder). Sucrose, fructose, and glucose elicited behavioral responses (proboscis extension reflex: PER) and electrophysiological responses (spikes) from ST. Sugar responsiveness was similar in both species, where sucrose was the most stimulatory. Two fermentation products, ethanol and acetic acid, never induced PERs but elicited large-amplitude spikes at a concentration of >1% (w/v). The two species significantly differed in responsiveness to the binary mixtures of sucrose and the fermentation products. Ethanol enhanced the sugar responses of V. indica but slightly inhibited those of A. hyperbius. Although acetic acid suppressed the sugar responses of both species, V. indica was less susceptible than A. hyperbius. When concentration of the fermentation products increased, binary mixtures evoked large spikes together with small ones regarded as the sucrose responses. Unlike the proboscal sensilla in our previous study, the tarsal ST of both species unambiguously responded to fermentation products. These results demonstrate that the tarsal gustatory sense of V. indica is adaptive to the use of rotting foods.     

Preferences for nectar sugars in the peacock butterfly, Inachis io

1. Peacock butterflies, Inachis io , were tested experimentally for their preferences for nectar sugars.
2. In tests with different plain sugar solutions (25%, weight to total weight) the butterflies strongly preferred sucrose and fructose over glucose. They also preferred sucrose over fructose.
3. In tests with mixed sugar solutions the butterflies clearly preferred both sucrose-dominant (sucrose : hexoses = 5 : 1) and balanced sugar solutions (sucrose : glucose : fructose = 1 : 1 : 1) over hexose-dominant sugar solutions (sucrose : hexoses = 1 : 5).
4. Females consumed significantly more of the balanced sugar solution than did males.
5. These results are discussed with respect to previous experiments on nectar preferences of butterflies, nectar sugar composition of butterfly-pollinated flowers, and flower preferences, physiological and reproductive aspects of butterflies.     

VARIATION IN NECTAR SUGAR COMPOSITION AT THE INTRAPLANT LEVEL IN IPOMOPSIS LONGIFLORA (POLEMONIACEAE)

The nectar sugar composition of individual flowers from single plants in both greenhouse and field populations was determined by High-Performance Liquid Chromatography (HPLC). Intraplant variability in nectar sugar composition of greenhouse-grown plants was significantly greater than can be attributed to the analytical methodology. Individual greenhouse-grown plants had significantly different sugar compositions, suggesting genetic differences. Under field conditions the situation was complex. Variation among flowers from single plants in field populations was significantly greater than that of plants under greenhouse conditions. Unprotected flowers of some field populations were more variable at the intraplant level than others. Field experiments showed that, in untreated (open-visited) flowers, the percent sucrose relative to other sugars declined significantly and coefficients of variability increased significantly with flower age. However, nectars never exposed to the natural environment did not change significantly with flower age. Further, within single plants it was found that percent sucrose in nectars varied in a small but significant way during the growing season. The data suggest that, in this species, both intrinsic and extrinsic factors affect nectar sugar composition and it is desirable to obtain samples from numerous flowers to accurately characterize the nectar sugar composition of an individual under field conditions.     

NECTAR PRODUCTION,COMPOSITION, ENERGETICS AND POLLINATOR ATTRACTIVENESS IN SPRING FLOWERS OF WESTERN NEW YORK

High Performance Liquid Chromatography (HPLC) was used to determine specific sugar ratios (fructose, glucose and sucrose) in nectars of nine families of flowering plants. All nectars contained all three sugars with the exception of that of Asclepias. Asclepias nectar was nearly a pure sucrose solution. Sucrose/hexose ratio was correlated with flower morphology, with tubular flowers having more sucrose and open flowers having more hexose. Open flowers contained nectar whose concentration was more affected by relative humidity than tubular flowers. Available nectar in unbagged flowers was found in volumes of 0.1 to 5 μl (17.5 to 68.2% sugar). Total sugar available per blossom amounted to 0.07 to 3.54 mg. We note that care must be taken in converting % concentration to mg sugar. Energetic yield was not as variable as the other measures and ranged from 0.72 to 3.58 cal/μl. Total daily nectar production was measured in five families and 24-hr sugar production varied from 0.64 to 5.52 mg per flower. Insect nectar feeders frequently searched many blossoms with little or no reward, but were rewarded sufficiently at rare “lucky hit” blossoms which contained relatively large nectar rewards. Insect pollinators did not seek nectars of specific sucrose-hexose ratios, but instead took nectar where caloric reward and accessibility made it most profitable.     

Nectar sugars enhance fitness in male Coenonympha pamphilus butterflies by increasing longevity or realized reproduction

The principal components of floral nectar are water and the sugars sucrose, fructose and glucose. Several studies have shown the importance of nectar sugars for female butterfly fecundity, whereas to date little attention has been paid to the effect of nectar sugars on male butterfly reproduction. Clear evidence for an effect of nectar sugars on male realized reproductive success is still missing. In this study, we fed male Coenonympha pamphilus butterflies nectar mimics with low (5%), medium (20%) or high (30%) total sugar concentrations with a sucrose:glucose:fructose ratio of 2.7:1.1:1. Sugar solutions were made mimicking Knautia arvensis, an essential nectar plant for C. pamphilus and many other European butterflies. Realized male reproductive success for each treatment was measured indirectly via nuptial gifts, by recording reproductive parameters and by characterizing time patterns over the oviposition period of their female partner. Male butterflies fed high‐concentrated nectar sugars had a longer lifespan than males fed low‐concentrated nectar sugars. In contrast, offspring of males fed medium‐concentrated nectar sugars had a higher hatching mass than progeny of males fed low‐concentrated nectar sugars, indicating a tradeoff between somatic maintenance and reproduction in the use of nectar sugars. Thus, allocation patterns of nectar sugars differed according to sugar concentrations in adult food. The method used in this experiment took into account the indispensable role of female butterflies in passing male nutrients to offspring. With this comprehensive approach, we can show the general importance of nectar sugars for male butterfly fitness and support previous findings suggesting a coevolutionary process between butterflies and flowers dependent on butterfly pollination.     

Scaling of nectar foraging in orchid bees

Morphology influences the rate at which foraging bees visit nectar flowers, the quantity of nectar they must consume to fuel their activities, and, consequently, the profitability of flower species. Because feeding time is a major determinant of visitation rate, I used a biomechanical model to examine how energy intake rate (E) varies with sucrose concentration, body mass (M), and proboscis length in orchid bees (Apidae: Euglossini). Under geometric scaling, the optimal sugar concentration (Smax) should be largely independent of body size, and E proportional to M1.0. In a comparative study of 30 orchid bee species ranging from 50 to 800 mg, Smax fell between 35% and 40% w/w, but E proportional to M0.54, significantly less than model predictions. Proboscis length and radius scale geometrically with body mass, but proboscis length exhibits substantial size-independent variation, particularly in small bees. One cost of a long proboscis is a reduction in both E and Smax in accordance with the scaling model. This finding highlights a difference between the lapping mechanism used by bumblebees and the suction mechanism used by orchid bees. A field study confirms that orchid bees harvest nectars with between 34% and 42% sucrose, independent of body size.     

Nectar to improve parasitoid fitness in biological control: Does the sucrose:hexose ratio matter?

The consumption of saccharide-rich foods such as floral nectar is crucial for the survival of many adult parasitoid wasps. The importance to parasitoids of nectar quality, with regards to its sucrose:hexose ratio, was investigated. Nectar, an aqueous solution of sugars, amino acids and other compounds, differs between plant species. Nectar composition is dominated by sucrose, glucose and fructose. Previous studies have shown that the ratio of sucrose to hexose (glucose+fructose) sugars can explain nectar associations in a range of flower visiting arthropods. It has been suggested that this ratio may be important in terms of parasitoid fitness. Analysis of floral nectar from fourteen plant species confirmed that the sucrose/hexose ratio significantly differed between species. An opportunity to select floral resources based on this measure of nectar quality arose and highlighted the potential to utilize native flowering plant species in place of the seven most commonly deployed, which are usually not native to the countries in which they are used.Results presented in this paper indicate, however, that the sucrose/hexose ratio is not a significant factor explaining parasitoid longevity. The hymenopteran parasitoids Diadegma semiclausum (Ichneumonidae) and Dolichogenidea tasmanica (Braconidae) were fed 40% w/w sugar solutions, differing in their sugar ratios. Solutions were classified as either sucrose-dominant (ratio >0.99), sucrose-rich (ratio 0.5–0.99), hexose-rich (ratio 0.1–0.499) or hexose-dominant (ratio <0.1). No significant differences in parasitoid longevity were found between the different treatments for either species. This suggests there is not an optimal sucrose/hexose ratio for parasitoid wasps, although a greater number of parasitoid species should ideally be tested to confirm if this is true for the wider parasitoid taxonomic groups.     

Floral nectaries, nectar production dynamics and chemical composition in six ipomoea species (convolvulaceae) in relation to pollinators

BACKGROUND AND AIMS: Floral nectaries and nectar features were compared between six Argentinian Ipomoea species with differences in their pollinator guilds: I. alba, I. rubriflora, I. cairica, I. hieronymi var. hieronymi, I. indica, and I. purpurea. METHODS: Pollinators were recorded in natural populations. The morpho-anatomical study was carried out through scanning electron and light microscopy. Nectar sugars were identified via gas chromatography. Nectar production and the effect of its removal on total nectar sugar amount were determined by using sets of bagged flowers. KEY RESULTS: Hymenopterans were visitors of most species, while hummingbirds visited I. rubriflora and sphingids I. alba. All the species had a vascularized discoidal nectary surrounding the ovary base with numerous open stomata with a species-specific distribution. All nectar samples contained amino acids and sugars. Most species had sucrose-dominant nectars. Flowers lasted a few hours. Mean nectar sugar concentration throughout the lifetime of the flower ranged from 34.28 to 39.42 %, except for I. cairica (49.25 %) and I. rubriflora (25.18 %). Ipomoea alba had the highest nectar volume secreted per flower (50.12 microL), while in the other taxa it ranged from 2.42 to 12.00 microL. Nectar secretion began as soon as the flowers opened and lasted for a few hours (in I. purpurea, I. rubriflora) or it was continuous during the lifetime of the flower (in the remaining species). There was an increase of total sugar production after removals in I. cairica, I. indica and I. purpurea, whereas in I. alba and I. rubriflora removals had no effect, and in I. hieronymi there was a decrease in total sugar production. CONCLUSIONS: The chemical composition, production dynamics and removal effects of nectar could not be related to the pollinator guild of these species. Flower length was correlated with nectary size and total volume of nectar secreted, suggesting that structural constraints may play a major role in the determination of nectar traits of these species.     

Pollinator type and secondarily climate are related to nectar sugar composition across the angiosperms

Pollinators are important agents of selection on floral traits, including nectar sugar composition. Although it is widely assumed that the proportion of sugars (mainly sucrose, glucose and fructose) in nectar reflects pollinators’ physiological limitations and digestive efficiency, the relative impact of pollinators and abiotic factors on nectar sugar composition, as well as the generality of these associations across the angiosperms, remain unknown. We compiled data on nectar sugar composition for >1000 plant species, along with information on flower visitors, plant growth form and latitudinal climatic zone, to provide the first comprehensive assessment of correlates of variation in sugar nectar composition in the angiosperms. After assembling a phylogeny linking all species in the dataset, we estimated the amount of phylogenetic signal in the percentage of sucrose and, by applying phylogenetically-informed multiple regressions, we evaluated whether nectar composition was influenced either by the main pollinator group, plant growth form, or latitudinal climatic zone. The relative importance of each of these factors was then assessed through model selection based on Akaike information criteria and deviance partitioning analysis. Nectar was dominated by sucrose in 56.8% of all the species, glucose in 16.7%, and fructose in 5.5%. Nectar in the remaining species was characterized by similar proportions of the three sugars. Variation in the proportion of sucrose was highest (~70%) at the intrafamily level, and had a significant but low phylogenetic signal, which partially reflects phylogenetic conservatism of the pollinator niche. After controlling for phylogenetic effects, the proportion of sucrose was mainly related to pollinator type and secondarily to climate. Accordingly, this study indicates that nectar sugar composition shows high evolutionary lability and its variation reflects plant-pollinator associations.     

Sugar Preferences in Nectar‐ and Fruit‐Eating Birds: Behavioral Patterns and Physiological Causes1

Sucrose, glucose, and fructose are the three sugars that commonly occur in floral nectar and fruit pulp. The relative proportions of these three sugars in nectar and fruit in relation to the sugar preferences of pollinators and seed dispersers have received considerable attention. Based on the research of Herbert and Irene Baker and their collaborators, a dichotomy between sucrose‐dominant hummingbird‐pollinated flowers and hexose‐dominant passerine flowers and fruits was proposed. Data on sugar preferences of several hummingbird species (which prefer sucrose) vs. a smaller sample of passerines (which prefer hexoses) neatly fitted this apparent dichotomy. This hummingbird–passerine dichotomy was strongly emphasized until the discovery of South African plants with sucrose‐dominant nectars, which are pollinated by passerines that are able to digest, and prefer sucrose. Now we know that, with the exception of two clades, most passerines are able to assimilate sucrose. Most sugar preference studies have been conducted using a single, relatively high, sugar concentration in the nectar (ca 20%). Thus, we lack information about the role that sugar concentration might play in sugar selection. Because many digestive traits are strongly affected not only by sugar composition, but also by sugar concentration, we suggest that preferences for different sugar compositions are concentration‐dependent. Indeed, recent studies on several unrelated nectar‐feeding birds have found a distinct switch from hexose preference at low concentrations to sucrose preference at higher concentrations. Finally, we present some hypotheses about the role that birds could have played in molding the sugar composition of plant rewards.     

Proboscis behavioral response of four honey bee Apis species towards different concentrations of sucrose,glucose, and fructose

Honey bees forage for pollen and nectar. Sugar is an important stimulus for foraging and a major source of energy for honey bees. Any differential response of bees to different concentrations of sugary nectar can affect their foraging. The sugar responsiveness of Apis species (Apis dorsata, Apis florea, and Apis cerana) was determined in comparison to that of Apis mellifera by evaluating the proboscis extension response (PER) with eight serial concentrations (0.00001, 0.0001, 0.001, 0.01, 0.1, 0.5, 1.0, and 1.5 M) of sucrose, glucose and fructose. Nectar foragers of bee species (A. dorsata, A. florea, A. cerana, and A. mellifera) exhibited an equal response for sucrose, glucose, and fructose, with no significant differences in their PER at all tested concentrations of these sugars within the same species. The inter-species comparison between Apis species revealed the differential responsiveness to the different concentrations of sugars, and the lowest concentration at which a response occurs was considered as the response threshold of these bee species for sugar solutions. A. mellifera presented significantly higher responsiveness than A. dorsata to low concentrations (0.00001, 0.0001, 0.001, 0.01, and 0.1 M) of sucrose, glucose and fructose. A. mellifera displayed a significantly higher response to water than A. dorsata. A. florea and A. mellifera presented no significant difference in their responsiveness to sucrose, glucose, and fructose at all tested concentrations, and their water responsiveness was also significantly at par but relatively higher in A. mellifera than in A. florea. Likewise, the responsiveness of A. cerana and A. mellifera to different concentrations of sucrose, glucose and fructose was significantly at par with no difference in their water responsiveness. This study represents preliminary research comparing the response of different honey bee species to three sugar types at different concentrations. The results imply that the native species are all better adapted than A. mellifera under local climate conditions.     

Feeding responses by female Pieris brassicae butterflies to carbohydrates and amino acids

Most Lepidoptera feed during the adult stage on carbohydrate‐rich food sources, primarily floral nectar. However, little is known about the factors leading to the acceptance of a possible food source. It is reported that butterflies select for nectar rich in sucrose and amino acids. This suggests that the insects have developed a sensitivity to these nectar compounds. We tested females of the large cabbage white, Pieris brassicae L. (Lepidoptera: Pieridae) for their responses to 10 different nectar‐ or honeydew‐sugars after either tarsal or proboscis stimulation. In no‐choice experiments, food‐deprived P. brassicae showed the strongest response to sucrose, followed by fructose. Other sugars, including galactose, glucose, maltose, mannose, melezitose, melibiose, raffinose and trehalose, did not elicit a feeding response. Mixtures of essential or common non‐essential amino acids did not stimulate feeding. In a choice situation, P. brassicae preferred sucrose over fructose, whereas they accepted a sucrose and amino acid solution equally to a plain sucrose solution. The results indicate that for P. brassicae, feeding is mainly elicited by sucrose and fructose.     

Field methods for sampling and storing nectar from flowers with low nectar volumes

Background and Aims

Although several methods of sampling and storing floral nectar are available, little information exists on sampling and storing nectar from flowers with low nectar volumes. Methods for sampling and storing nectar from the flowers of species with low floral nectar volumes (<1 µL) were investigated using the flowers of Eucalyptus species.

Methods

Sampling with microcapillary tubes, blotting up with filter paper, washing and rinsing were compared to determine masses of sugars recovered and differences in sugar ratios. Storage methods included room temperature, refrigeration and freezing treatments; the addition of antimicrobial agents benzyl alcohol or methanol to some of these treatments was also evaluated. Nectar samples were analysed using high-performance liquid chromatography and the masses of sucrose, glucose and fructose in each sample were determined.

Key Results

Masses of sugars varied significantly among sampling treatments, but the highest yielding methods, rinsing and washing, were not significantly different. A washing time of 1 min was as effective as one of 20 min. Storage trials showed that the sugar concentration measurements of nectar solutions changed rapidly, with the best results achieved for refrigeration with no additive (sucrose and fructose were stable for at least 2 weeks). Sugar ratios, however, remained relatively stable in most treatments and did not change significantly across 4 weeks for the methanol plus refrigerator and freezing treatments, and 2 weeks for the refrigeration treatment with no additive.

Conclusions

Washing is recommended for nectar collection from flowers with low nectar volumes in the field (with the understanding that one wash underestimates the amounts of sugars present in a flower), as is immediate analysis of sugar mass. In view of the great variation in results depending on nectar collection and storage methods, caution should be exercised in their choice, and their accuracy should be evaluated. The use of pulsed amperometric detection, more specific than refractive index detection, may improve the accuracy of nectar sugar analysis.Key words: Eucalyptus, flower with small nectar volume, nectar collection, nectar sampling, nectar storage, sugar ratio     

Host-plant shifts and adult survival in the cactus breeding Drosophila mojavensis

Abstract. 1. In order to gain insight into how Drosophila mojavensis Patterson and Crow (which lives on the rotting tissue of cactus) successfully shifted host plants, from an Opuntia species to columnar cactus species, over evolutionary time, specific components of columnar cactus and Optunia tussue were examined for their effects on adult longevity.
2. The availability of free sugars affects the survival of adult D.mojavensis . The addition of a sugar to columnar cactus tissue significantly increased the longevity of adults that fed on those tissues. This increase in adult longevity was equal to the longevity observed on unsupplemented Opuntia tissue, which has free sugars present.
3. Live microorganisms added to Opuntia tissue had a slight detrimental effect on adult longevity (longevity was decreased by approximately 1 week).
4. Low concentrations of atmospheric ethanol (independent of diet) increased the longevity of adult D.mojavensis to levels comparable in magnitude to that provided by the addition of free sugars to the diet.
5. This information and the apparent lack of a year-round supply of free sugars for D.mojavensis in nature suggests that the ethanol (produced by fermentation) rather than free sugars per se is the most important component of rotting cactus tissue for the survival of this species and may explain its successful host-plant shift.     

Floral nectary structure and nectar composition in Eccremocarpus scaber (Bignoniaceae), a hummingbird-pollinated plant of central Chile

Surface features, anatomy, and ultrastructure of the floral nectary of Eccremocarpus scaber (Bignoniaceae), pollinated predominantly by the largest-known hummingbird (Patagona gigas gigas), were studied together with nectar sugar content and secretion rate. The annular disk nectary comprises epidermis, secretory and ground parenchyma with intercellular spaces, and branched vascular bundles terminating in the secretory parenchyma where only phloem is found. Amyloplasts and vacuoles increase in size throughout development, the latter becoming sites of organelle degradation. Transferlike cells in nectary phloem and P-proteinlike fibrillar material in phloem parenchyma were observed. Flowers produced around 32 μl of nectar (mostly after anthesis) with 11 mg of sugar composed of fructose, glucose, sucrose, and maltose in a ratio of 0.34:0.32:0.17:0.17. Morphological studies as well as the presence of maltose and glucose in nectar suggest storage of the originally phloem-derived sugars as starch with its subsequent hydrolysis. The low sucrose/hexose ratio (0.25) and high nectary secretion force (nectar per flower biomass) observed places E. scaber close to large-bodied bat-pollinated plants. A hypothesis based on nectar origin and nectar secretion is advanced to explain pollinator-correlated variation in sucrose/hexose ratio.     

Extreme intraplant variation in nectar sugar composition in an insect-pollinated perennial herb

Variation in nectar chemistry among plants, flowers, or individual nectaries of a given species has been only rarely explored, yet it is an essential aspect to our understanding of how pollinator-mediated selection might act on nectar traits. This paper describes variation in nectar sugar composition in a population of the perennial herb Helleborus foetidus (Ranunculaceae) and dissects it into components due to variation among plants, flowers of the same plant, and nectaries of the same flower. The proportions of sucrose, glucose, and fructose in single-nectary nectar samples collected at two times in the flowering season were determined using high performance liquid chromatography (HPLC). Sugar composition varied extensively among nectaries, and nearly all combinations of individual sugars were recorded. Population-wide variance was mainly accounted for by variation among flowers of the same plant (56% of total), nectaries of the same flower (30%), and only minimally by differences among plants (14%). In absolute terms, intraplant variation was similar to or greater than that ordinarily reported in interspecific comparisons. Results suggest that the prevailing notion of intraspecific constancy in nectar sugar composition may be unwarranted for some species and that more elaborate nectar sampling designs are required to detect and appropriately account for extensive within-plant variance. Within-plant variation in nectar sugar composition will limit the ability of pollinators to exert selection on nectar chemistry in H. foetidus and may be advantageous to plants by reducing the number of flowers visited per foraging bout by variance-sensitive, risk-averse pollinators.     

Nectar sugar preferences and absorption in a generalist African frugivore, the Cape White-eye Zosterops pallidus

Sugar preferences and absorption efficiencies were investigated in the Cape White-eye Zosterops pallidus , a generalist frugivore. Unlike the mainly American frugivorous passerines previously studied, Cape White-eyes preferred 20% (weight: weight) solutions of sucrose to glucose or fructose, and apparently possess high sucrase activity because absorption efficiencies for all three sugars were close to 100%. Cape White-eyes rejected 20% solutions containing a fourth nectar sugar, xylose, recently found in the nectar of Protea and Faurea (Proteaceae). Absorption efficiencies for xylose averaged 61%, but we do not know whether this sugar is utilized by the birds. Cape White-eyes thus resemble specialized southern African nectarivores (Nectariniidae, Promeropidae) in their responses to nectar sugars.     

The effect of genotype, age, sex, and caste on response thresholds to sucrose and foraging behavior of honey bees (Apis mellifera L.)

Bees derived from artificially selected high- and low-pollen-hoarding strains were tested for their proboscis extension reflex response to water and varying sucrose concentrations. High-strain bees had a lower response threshold to sucrose than low-strain bees among pre-foragers, foragers, queens and drones. Pre-foraging low-strain workers showed ontogenetic changes in their response threshold to sucrose which was inversely related to age. High-strain foragers were more likely to return with loads of water compared to low-strain foragers. Whereas low-strain foragers were more likely to return with loads of nectar. Low-strain nectar foragers collected nectar with significantly higher sucrose concentrations than did the high-strain nectar foragers. Alternatively, low-strain foragers were more likely to return empty compared to high-strain foragers. These studies demonstrate how a genotypically varied sensory-physiological process, the perception of sucrose, are associated with a division of labor for foraging. Accepted: 27 October 1998     

FLORAL NECTAR SUGAR COMPOSITION AND POLLINATOR TYPE AMONG NEW WORLD GENERA IN TRIBE ANTIRRHINEAE (SCROPHULARIACEAE)

The floral nectar sugar compositions of 20 New World species from 10 genera and of five interspecific hybrids in tribe Antirrhineae have been analyzed using high-performance liquid chromatography. Species are pollinated by short-tongued bees, long-tongued bees, and hummingbirds. Ornithophily represents the derived condition in the tribe and has arisen independently in subtribes Maurandyinae and Gambeliinae. All nectars analyzed are sucrose-dominant or -rich, except for the hexose-rich nectar of Mohavea breviflora. Despite the predominance of sucrose, floral nectar sugars from species pollinated by different pollen vectors have characteristic constituents. Nectar sugars from flowers visited by hummingbirds average 76.2% sucrose and have compositions remarkably similar to hummingbird nectars analyzed in previous studies of unrelated species. Long-tongued bee nectars average 87% sucrose and differ from shorttongued bee nectars which have the lowest mean sucrose percentage (40.2%). The association of sugar constituent types and principal pollinators is concordant with previous data and supports hypotheses concerning pollinator preferences and the adaptive significance of certain nectar sugar compositions. Within this adaptive framework, phylogenetic constraint is also operative and may explain the predominance of sucrose in nectar sugars, similarities in sugar composition among hummingbird nectars in subtribes Maurandyinae and Gambeliinae, and the similarity of nectar from Galvezia leucantha (long-tongued bee-pollinated) to hummingbird-pollinated species also in subtribe Gambeliinae.