Ineffectiveness of nectar scent in generating bumblebee visits to flowers of Impatiens textori

To clarify if bumblebees can recognize nectar through its scent in Impatiens textori flowers, we examined the behavior of Bombus diversus on nectarless flowers in which the spurs had been artificially removed. Bumblebee visits to both natural flowers and spur‐cut flowers were captured using a long‐term video recording system. Visiting behavior and frequency were compared between the two flower types. Many bumblebees visited both types of flower, and their visit frequencies were not significantly different. However, the length of stay on each flower type did differ, with the bumblebees remaining on the spur‐cut flowers for a significantly shorter time than on the natural flowers. Our results suggest that bumblebees cannot detect the absence of nectar in I. textori flowers before probing them. Therefore, the nectar scent of I. textori does not serve to attract bumblebees although the presence of nectar will detain bumblebees on flowers for longer periods.     

Discrimination of Unrewarding Flowers by Bees; Direct Detection of Rewards and Use of Repellent Scent Marks

Bumblebees and honeybees deposit short-lived scent marks on flowers that they visit when foraging. Conspecifics use these marks to distinguish those flowers that have recently been emptied and, so, avoid them. The aim of this study was to assess how widespread this behavior is. Evidence for direct detection of reward levels was found in two bee species: Agapostemon nasutus was able to detect directly pollen availability in flowers with exposed anthers, while Apis mellifera appeared to be able to detect nectar levels of tubular flowers. A third species, Trigona fulviventris, avoided flowers that had recently been visited by conspecifies, regardless of reward levels, probably by using scent marks. Three further bee/flower systems were examined in which there was no detectable discrimination among flowers. We argue that bees probably rely on direct detection of rewards where this is allowed by the structure of the flower and on scent marks when feeding on flowers where the rewards are hidden. However, discrimination does not always occur. We suggest that discrimination may not always make economic sense; when visiting flowers with a low handling time, or flowers that are scarce, it may be more efficient to visit every flower that is encountered.     

Studies on ecology and behavior of Japanese black swallowtail butterflies. 6. Nectar feeding ofPapilio helenus nicconicolens Butler andP. protenor demetrius Cramer as main pollinators of glory bower,Clerodendron trichotomum,Thunb

The butterfliesPapilio helenus andP. protenor were shown to feed mainly on the nectar of the glory bower,Clerodendron trichotomum, which was the most abundant nectar plant in summer in the study area. Both the species were found to have a proboscis longer than 24 mm corresponding to the length of the corolla tube ofC. trichotomum. Visits to the flowers by these butterflies were observed more frequently than visits by sphingid moths which had previously been believed to be the major pollinators ofC. trichotomum. The male butterflies visited trees ofC. trichotomum frequently, while visits by the females were less frequent. However, once females had visited the tree ofC. trichotomum, they remained there longer than the males. Since the flower ofC. trichotomum has long protruding sexual organs, its pollen grains were found to adhere efficiently to the bodies of butterflies, mainly the thorax, during nectar feeding. Most of the butterflies became loaded withC. trichotomum pollen, and the mean number of pollen grains per butterfly was 1,776 forP. helenus and 2,817 forP. protenor. The flowers opened at any time of day but more frequently in the morning. The nectar was secreted throughout the day. In the maturation of the protandrous flower ofC. trichotomum, the duration of the pistillate phase was about twice as long as the staminate phase. The long flowering period and the short duration of the staminate phase resulted in asynchrony of the flowering stages even within a single cyme on a tree. Such asynchrony and the abundance of attractive flowers on a tree facilitates efficient pollination by the butterflies.     

Why do nectar-foraging bees and wasps work upwards on inflorescences?

Summary Wasps (Dolichovespula and Vespula spp.) worked predominantly upwards when foraging for nectar on inflorescences of the protogynous Scrophularia aquatica, in which the standing crop of nectar sugar per flower showed no clear pattern of vertical distribution up an inflorescence. Bumblebees taking nectar (Bombus hortorum visiting legally, and certain individuals of B. terrestris which positioned themselves head-upwards while taking nectar through holes bitten in the corolla) worked predominantly upwards on the racemose inflorescences of Linaria vulgaris, although the standing crop of nectar sugar per open flower increased up the raceme. Individuals of B. terrestris which robbed Linaria flowers in a head-down position worked predominantly downwards on inflorescences. The upward or downward directionality of intra-inflorescence movements by foraging insects may depend in part on the position these adopt during their flower visits.     

Specialised protagonists in a plant–pollinator interaction: the pollination of Blumenbachia insignis (Loasaceae)

• Analyses of resource presentation, floral morphology and pollinator behaviour are essential for understanding specialised plant‐pollinator systems. We investigated whether foraging by individual bee pollinators fits the floral morphology and functioning of Blumenbachia insignis, whose flowers are characterised by a nectar scale‐staminode complex and pollen release by thigmonastic stamen movements.
• We described pollen and nectar presentation, analysed the breeding system and the foraging strategy of bee pollinators. We determined the nectar production pattern and documented variations in the longevity of floral phases and stigmatic pollen loads of pollinator‐visited and unvisited flowers.
• Bicolletes indigoticus (Colletidae) was the sole pollinator with females revisiting flowers in staminate and pistillate phases at short intervals, guaranteeing cross‐pollen flow. Nectar stored in the nectar scale‐staminode complex had a high sugar concentration and was produced continuously in minute amounts (~0.09 μl·h^?1). Pushing the scales outward, bees took up nectar, triggering stamen movements and accelerating pollen presentation. Experimental simulation of this nectar uptake increased the number of moved stamens per hour by a factor of four. Flowers visited by pollinators received six‐fold more pollen on the stigma than unvisited flowers, had shortened staminate and pistillate phases and increased fruit and seed set.
• Flower handling and foraging by Bicolletes indigoticus were consonant with the complex flower morphology and functioning of Blumenbachia insignis. Continuous nectar production in minute quantities but at high sugar concentration influences the pollen foraging of the bees. Partitioning of resources lead to absolute flower fidelity and stereotyped foraging behaviour by the sole effective oligolectic bee pollinator.

     

Morphological variation in relation to flower use in bumblebees

To understand resource partitioning in a bumblebee community, we analyzed various morphological characters. A total of 1269 individuals of six bumblebee species, Bombus ardens, B. hypocrita, B. diversus, B. ignitus, B. honshuensis and B. beaticola, were examined and principal component analysis showed that the bumblebee species were clearly differentiated. Glossa, prementa and head lengths were positively correlated with the second component, and a longer proboscis was associated with a narrower body, which may help bees to intrude into and access deep‐lying nectar sources. Bombus diversus, with a long proboscis and narrow body, preferred flowers with a long corolla tube, whereas B. hypocrita and B. ignitus, which have short proboscises and wide bodies, visited flowers with short corollas or dish‐shaped flowers. Two pairs of consubgeneric species that have similar morphological characteristics, B. ardens and B. beaticola, and B. hypocrita and B. ignitus, divided flower resources by habitat selection and seasonal partitioning. For resource partitioning among bumblebee species, not only morphology but also other factors, such as habitat and seasonal preference, flower use, foraging behavior, and interspecific interactions, are responsible.     

Bumblebee-pollination and temporal change of the calyx tube length in Clematis stans(Ranunculaceae)

Clematis stans is dioecious semi-arboreal, with pale purple–blue, nodding, tubulous flowers in a paniculate inflorescence. Both male and female flowers produce nectar from the base of the calyx tube during a flowering period of 3 or 4 days, and are pollinated by two bumblebee species, Bombus diversus and B. honshuensis, with different proboscis lengths. When the flowers open, four sepals constructing a calyx tube separate at the top and their respective tips gradually curl up, so that a tubular part shortens. Observations at two field sites showed that B. diversus (with a longer proboscis) most often visits the flowers with a longer calyx tube, and B. honshuensis (with a shorter proboscis) the flowers with a shorter calyx tube, i.e., later in the flowering period. By changing the calyx tube length, the flowers of C. stans accept the two bumblebee species with different proboscis length as pollinators and thus increase the chance of pollination for each flower. It was also found that the two bumblebee species prefer the male flowers to the female flowers, although the female flowers secrete more nectar as a reward than male flowers. This is likely because they visit the male flowers to collect pollen grains in addition to nectar. Electronic Publication     

Bumblebee Pollination of Melampyrum ciliare (Scrophulariaceae)

Abstract Melampyrum ciliare has tubular flowers predominantly visited by Bombus diversus. Floral structure (including the positions of anthers and stigma) and structure of the distal part of the corolla indicate cross-pollination by bumblebees. In M. ciliare, young flowers with white spots on the labium, which disappeared with flower aging, produced larger amounts of nectar than older ones without spots. Bumblebees visited flowers with white spots significantly more frequently than would be expected if they chose flowers randomly. These findings and the high seed production of open-pollinated flowers suggest effective pollination of M. ciliare by bumblebees.     

Intra‐floral resource partitioning between endemic and invasive flower visitors: consequences for pollinator effectiveness

1. Sympatric flower visitor species often partition nectar and pollen and thus affect each other's foraging pattern. Consequently, their pollination service may also be influenced by the presence of other flower visiting species. Ants are solely interested in nectar and frequent flower visitors of some plant species but usually provide no pollination service. Obligate flower visitors such as bees depend on both nectar and pollen and are often more effective pollinators. 2. In Hawaii, we studied the complex interactions between flowers of the endemic tree Metrosideros polymorpha (Myrtaceae) and both, endemic and introduced flower‐visiting insects. The former main‐pollinators of M. polymorpha were birds, which, however, became rare. We evaluated the pollinator effectiveness of endemic and invasive bees and whether it is affected by the type of resource collected and the presence of ants on flowers. 3. Ants were dominant nectar‐consumers that mostly depleted the nectar of visited inflorescences. Accordingly, the visitation frequency, duration, and consequently the pollinator effectiveness of nectar‐foraging honeybees (Apis mellifera) strongly decreased on ant‐visited flowers, whereas pollen‐collecting bees remained largely unaffected by ants. Overall, endemic bees (Hylaeus spp.) were ineffective pollinators. 4. The average net effect of ants on pollination of M. polymorpha was neutral, corresponding to a similar fruit set of ant‐visited and ant‐free inflorescences. 5. Our results suggest that invasive social hymenopterans that often have negative impacts on the Hawaiian flora and fauna may occasionally provide neutral (ants) or even beneficial net effects (honeybees), especially in the absence of native birds.     

The movement patterns of carpenter beesXylocopa micans and bumblebeesBombus pennsylvanicus onPontederia cordata inflorescences

The movement patterns of carpenter bees (Xylocopa micans) and bumblebees (Bombus pennsylvanicus) foraging for nectar on vertical inflorescences ofPontederia cordata were studied near Miami, Florida. The floral biology ofP. cordata is unique in several ways: (a) many short-lived flowers per inflorescence, (b) constant nectar production throughout the life span of each flower, and (c) abscence of vertical patterning of nectar and age of flowers. Inflorescences ranged between 3.5 and 15.8 cm long and had between 9 and 55 open flowers. Both carpenter bees and bumblebees arrived mostly on the bottom third of the inflorescence and left after visiting flowers on the top third of the inflorescence. The departure position from the inflorescence was higher up than observed in studies of other insect pollinators foraging on other speces of plants. This pattern of departure probably occurs in the absence of a vertical gradient of nectar or floral morphology.     

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

Abstract.

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

     

Nectar secretion pattern of the dish-shaped flower,Cayratia japonica (Vitaceae), and nectar utilization patterns by insect visitors

We studied the relationship between the diurnal nectar secretion pattern of flowers of Cayratia japonica and insect visiting patterns to these flowers. Flower morphology of C. japonica changed greatly for about 12 hours after flower-opening and the maximum duration of nectar secretion was 2 days. The nectar volume peaked at 11∶00 and 15∶00, and declined at night and at 13∶00 regardless of time elapsed after flower-opening. The nectar volume at the two peaks was, on average, 0.25 μl on bagged inflorescences and 0.1μl on unbagged inflorescences (both, sugar concentration=60%). The flower secreted nectar compensatory when the nectar was removed. This means that insects consume more nectar than the difference of nectar volume between bagged and unbagged flowers. Apis cerana is a primary visitor of this flower, and was the only species for which we confirmed pollen on the body, among many species of flower visiting insects to this flower. Apis cerana visited intensively at the two peaks of nectar secretion. Visits of the other insects were rather constant or intensive only when there was no nectar secretion. Thus flowers of C. japonica with morphologically unprotected nectaries may increase likelihood that their nectar is used by certain pollinators, by controlling the nectar secretion time in day. In this study the pattern of nectar secretion allowed A. cerana maximum harvest of nectar.     

Male-biased sex ratio and promiscuous pollination in the dioecious island treeLaurus azorica (Lauraceae)

Pollination ofLaurus azorica (Lauraceae), a dioecious Macaronesian tree, was studied. Male and female trees had the same size distribution. The population had 2.5 times as many male trees as females. In addition, males produced more flowers, and their inflorescences lasted longer. Individual flower lifetime and length of flowering season were the same in both sexes. Between the years of observation, one tree changed sex. Pollinators wereHalictinae bees and the flyTachina canariensis. The bees collected pollen and nectar and the fly collected nectar from both sexes. Both species visited other plants as well. The evolution of breeding systems inLauraceae is discussed.     

Interference and exploitation competition of three nectar-thieving invasive ant species

Summary. Plant and insect exudates are known to play a key role in structuring tropical ant communities, but less is known about the utilization of these resources in communities dominated by invasive ants. Invasive ants are thought to require large amounts of carbohydrates such as honeydew or nectar to maintain their high abundances. Invasive ants that consume floral nectar may compete with legitimate floral visitors through interference or exploitation competition. I compared the nectar-thieving behavior of three widespread invasive ant species: long-legged ants (Anoplolepis gracilipes), Argentine ants (Linepithema humile), and big-headed ants (Pheidole megacephala) in inflorescences of the native Hawaiian ‘ōhi’a tree, an important food source for native fauna. A. gracilipes was least likely to leave inflorescences unvisited and visited inflorescences in higher numbers than both L. humile and P. megacephala. A. gracilipes and L. humile visited more flowers in an inflorescence and were less likely to retreat from a flower with a competitor than P. megacephala. A. gracilipes was able to take 5.5 and 11.3 times the amount of nectar than L. humile and P. megacephala, respectively. Thus, A. gracilipes may be effective at both interference and exploitation competition against other nectarivores, L. humile may be effective at interference competition, and P. megacephala may be relatively weak at both types of competition against other nectarivores. Ascertaining the competitive abilities of invasive ants against legitimate floral visitors will be especially important in agricultural and other systems that are nectar or pollinator limited.Received 6 December 2004; revised 13 January 2005; accepted 14 January 2005.     

Mobility of Impatiens capensis flowers: effect on pollen deposition and hummingbird foraging

Flexible pedicels are characteristic of birdpollinated plants, yet have received little attention in studies of hummingbird-flower interactions. A major implication of flexible pedicels is that flowers may move during pollination. We examined whether such motion affected interactions between ruby-throated hummingbirds (Archilochus colubris) and jewelweed (Impatiens capensis) by increasing pollen deposition and by altering the effectiveness of nectar removal. For I. capensis, flower mobility enhanced pollen deposition: birds had significantly longer contact with anthers and more pollen deposited on their bills and crowns when foraging at mobile flowers than at flowers that had been experimentally immobilized. In contrast, flower mobility imposed a cost on hummingbirds by significantly increasing their handling times and reducing their extraction rates relative to their interactions with immobile flowers. Field observations indicated that the motion observed during hummingbird visits did not occur when bees (Bombus spp., Apis mellifera) visited I. capensis flowers, which suggests that the mobility of I. capensis flowers is an adaptation for hummingbird pollination.     

The learning abilities of the white cabbage butterfly,Pieris rapae,foraging for flowers

This study examines the role of learning and memory in the butterflyPieris rapae crucivora Boisduval during foraging for flowers. In an outdoor cage with 6 flower species,P. rapae showed various visiting patterns: some visited only one species, while others visited several species in a day. The foraging process for flowers ofErigeron annuus (L.) Pers. could be divided into two successive steps: (1) landing on the nectaring caputs, and (2) finding the source of nectar in the caput. Butterflies learned to proceed through the two steps more efficiently with successive attempts: they gradually decreased landings on nectarless caputs and probings on the nectarless petals of ligulate flowers respectively. As a result, handling time per unit caputs became shorter, and apparent rewards per unit time, i.e. the efficiency of collecting nectar, increased. In addition, once learned,P. rapae could remember a rewarding flower color for 3 days, which was not interfered with by learning another flower color. This indicates thatP. rapae keeps memory for a period longer than 3 days, and that they can remember at least two flower species as suitable flower resources. Furthermore, data indicated that they sometimes can apply the foraging skills obtained on other flower species to a novel one. These abilities could enable butterflies to easily switch flower species, or to enhance labile preference. It has been known thatP. rapae also shows flower constancy, which may be due to memory constraints. Therefore, they may appropriately use two foraging tactics: visit consistency and labile preference, to get enough nectar according to their circumstances.     

Effects of Foraging Experiences on Residence Time of the Predatory Mite <Emphasis Type="BoldItalic">Neoseiulus womersleyi</Emphasis> in a Prey Patch

We investigated the effects of foraging experiences on the residence time of Neoseiulus womersleyi in a currently inhabited prey (Tetranychus urticae) patch. Satiated predators that had experienced starvation stayed longer in a current patch than those that had not experienced starvation. Satiated predators that had experienced a prey-rich patch showed approximately the same residence time in the current patch irrespective of the number of prey therein. By contrast, satiated predators that had experienced a prey-poor patch stayed longer in a current patch of high prey density than in one of low prey density. N. womersleyi appears to determine residence time in the current patch based on foraging experiences together with the quantity of prey in the current patch.     

Choice of flowers by foraging honey bees (Apis mellifera): possible morphological cues

Abstract.

• 1 Honey bees foraging for nectar on lavender (Lavandula stoechas) chose inflorescences with more of their flowers open. The number of open flowers predicted whether an inflorescence was visited by bees, inspected but rejected, or ignored. Inflorescences chosen arbitrarily by observers had numbers of open flowers intermediate between those of visited and ignored inflorescences.
• 2 Differences in morphological characters between types of inflorescence correlated with nectar volume and sugar weight per flower so that visited inflorescences had a disproportionately greater volume of nectar and weight of sugar per flower and greater variance in nectar volume.
• 3 Although there were significant associations between nectar content and the morphological characters of inflorescences, discriminant function analysis revealed discrimination on the basis of morphology rather than nectar content.
• 4 Visited inflorescences tended to have smaller than average flowers but bees tended to probe the largest flowers on visited inflorescences.
• 5 Choice of flowers within inflorescences is explicable in terms of the relationship between flower size and nectar content.

     

Patterns of plant visitation by nectar-feeding lizards

Geckos in the genus Hoplodactylus visit flowers to feed on nectar. I examined the patterns of flower visitation exhibited by two gecko species (H. maculatus and H. duvauceli) having access to two plant species: pohutukawa (Metrosideros excelsa: Myrtaceae) and flax (Phormium tenax: Agavaceae). Individual geckos were not observed to visit both plant species; individuals visiting flax tended to revisit the same plant. Geckos visiting pohutukawa were larger than those visiting flax and exhibited an early night peak in plant visitation, while lizards on flax displayed a more even pattern of activity throughout the night. On flax, geckos were more likely to be found on plants with a greater number of male flowers. Male flax flowers were of greater diameter than female flowers and produced nectar at higher rates and with greater concentrations of sugars. Experimental manipulation of pohutukawa nectar volumes suggested that the distribution of geckos is influenced by the pattern of nectar availability.     

Pollen foraging by bumblebees: Foraging patterns and efficiency on Lupinus polyphyllus

Summary Bumblebees foraging on vertical inflorescences start near the bottom and work upward, behavior commonly interpreted as a response to the greater amounts of nectar available in lower flowers. Lupinus polyphyllus, which produces no nectar, has more pollen available in upper flowers. Although bees are probably unable to detect this gradient, since pollen is hidden from their view, they still start low and forage upward. Therefore, we concluded that the bees' tendency to forage upward on vertical inflorescences is not tied to a reward gradient. In addition, bees use only about 15% of the flowers per inflorescence, although they could be much more efficient by visiting and revisiting every flower systematically. In general, revisits would not be penalized because most flowers contain enough pollen for several visits. Optimal foraging theory may not offer an adequate explanation for such gross inefficiency.