Behavioural differences between male and female carpenter bees in nectar robbing and its effect on reproductive success in Glechoma longituba (Lamiaceae)

Male and female nectar robbers may show significantly different behaviour on host plants and thus have different impacts on reproductive fitness of the plants. A 4-year study in natural populations of Glechoma longituba has shown that male carpenter bees (Xylocopa sinensis) are responsible for most of the nectar robbing from these flowers, while female bees account for little nectar robbing, demonstrating distinct behavioural differentiation between male and female bees in visiting flowers. The smaller male bee spends less time visiting a single flower than the larger female bee, consequently, the male bee is capable of visiting more flowers per unit time and has a higher foraging efficiency. Moreover, the robbing behaviour of female carpenter bees is more destructive and affects flower structures (ovules and nectaries) and floral life-span more than that of the male bee. According to the energy trade-off hypothesis, the net energy gain for male bees during nectar robbing greatly surpasses energy payout (17.72 versus 2.43 J), while the female bee net energy gain is barely adequate to meet energy payout per unit time (3.78 versus 2.39 J). The differences in net energy gain for male and female bees per unit time in nectar robbing are the likely cause of observed behavioural differences between the sexes. The differences in food resource preference between male and female bees constitute an optimal resource allocation pattern that enables the visitors to utilise floral resources more efficiently.     

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.

     

Nectar robbery by bees Xylocopa virginica and Apis mellifera contributes to the pollination of rabbiteye blueberry

Honey bees, Apis mellifera L., probe for nectar from robbery slits previously made by male carpenter bees, Xylocopa virginica (L.), at the flowers of rabbiteye blueberry, Vaccinium ashei Reade. This relationship between primary nectar robbers (carpenter bees) and secondary nectar thieves (honey bees) is poorly understood but seemingly unfavorable for V. ashei pollination. We designed two studies to measure the impact of nectar robbers on V. ashei pollination. First, counting the amount of pollen on stigmas (stigmatic pollen loading) showed that nectar robbers delivered fewer blueberry tetrads per stigma after single floral visits than did our benchmark pollinator, the southeastern blueberry bee, Habropoda laboriosa (F.), a recognized effective pollinator of blueberries. Increasing numbers of floral visits by carpenter bee and honey bee robbers yielded larger stigmatic loads. As few as three robbery visits were equivalent to one legitimate visit by a pollen-collecting H. laboriosa female. More than three robbery visits per flower slightly depressed stigmatic pollen loads. In our second study, a survey of 10 commercial blueberry farms demonstrated that corolla slitting by carpenter bees (i.e., robbery) has no appreciable affect on overall V. ashei fruit set. Our observations demonstrate male carpenter bees are benign or even potentially beneficial floral visitors of V ashei. Their robbery of blueberry flowers in the southeast may attract more honey bee pollinators to the crop.     

Floral Temperature and Optimal Foraging: Is Heat a Feasible Floral Reward for Pollinators?

As well as nutritional rewards, some plants also reward ectothermic pollinators with warmth. Bumble bees have some control over their temperature, but have been shown to forage at warmer flowers when given a choice, suggesting that there is some advantage to them of foraging at warm flowers (such as reducing the energy required to raise their body to flight temperature before leaving the flower). We describe a model that considers how a heat reward affects the foraging behaviour in a thermogenic central-place forager (such as a bumble bee). We show that although the pollinator should spend a longer time on individual flowers if they are warm, the increase in total visit time is likely to be small. The pollinator's net rate of energy gain will be increased by landing on warmer flowers. Therefore, if a plant provides a heat reward, it could reduce the amount of nectar it produces, whilst still providing its pollinator with the same net rate of gain. We suggest how heat rewards may link with plant life history strategies.     

Remote perception of floral nectar by bumblebees

Summary On both artificial flowers in the laboratory and certain plant species in the field, bumblebees often closely approached flowers and then departed without probing for nectar. In laboratory experiments where nectar rewards were associated with subtle visual or olfactory cues, bumblebees approached and avoided non-rewarding flowers. Flowers that bees entered and probed for nectar contained rewards much more frequently than predicted by chance alone. When there were no external cues associated with nectar content, bees visited rewarding flowers by chance alone, provided rewarding flowers were not spatially clumped. In the field, bumblebees approached and rejected a large proportion of dogbane flowers and red clover inflorescences. On both species, flowers or inflorescences probed by bees contained more nectar than those rejected by bees or those that I chose at random. On fireweed and monkshood, bees rarely or never approached and rejected healthy-looking flowers. Predictions generated by an optimal foraging model were tested on data from four bumblebee species foraging on red clover. The model was highly successful in qualitatively predicting the relationship between handling time and proportion of inflorescences rejected by individual bees, and the relationship between threshold nectar content for acceptance by bees and average resource availability. Thus, bees appeared to use remotely perceived cues to maximize their rates of nectar intake.     

Reversal Learning and Risk‐Averse Foraging Behavior in the Monarch Butterfly,Danaus plexippus (Lepidoptera: Nymphalidae)

Learning ability allows insects to respond to a variable environment, and to adjust their behaviors in response to positive or negative experiences. Pollinating insects readily learn to associate floral characteristics, such as color, shape, or pattern, with appetitive stimuli, such as the presence of a nectar reward. However, in nature pollinators may also encounter flowers that contain distasteful or toxic nectar, or offer highly variable nectar volumes, providing opportunities for aversive learning or risk‐averse foraging behavior. Whereas some bees learn to avoid flowers with unpalatable or unreliable nectar rewards, little is known about how Lepidoptera respond to such stimuli. We used a reversal learning paradigm to establish that monarch butterflies learn to discriminate against colored artificial flowers that contain salt solution, decreasing both number of probes and probing time on flowers of a preferred color and altogether avoiding artificial flowers of a non‐preferred color. In addition, when we offered butterflies artificial flowers of two different colors, both of which contained the same mean nectar volume but which differed in variance, the monarchs exhibited risk‐averse foraging: they probed the constant flowers significantly more than the variable ones, regardless of flower color or butterfly sex. Our results add to our understanding of butterfly foraging behavior, as they demonstrate that monarchs can respond to aversive as well as appetitive stimuli, and can also adjust their foraging behavior to avoid floral resources with high variance rewards.     

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.     

Pollination in Jacaranda rugosa (Bignoniaceae): euglossine pollinators, nectar robbers and low fruit set

Nectar robbers access floral nectar in illegitimate flower visits without, in general, performing a pollination service. Nevertheless, their effect on fruit set can be indirectly positive if the nectar removal causes an incremental increase in the frequency of legitimate flower visits of effective pollinators, especially in obligate outcrossers. We studied pollination and the effect of nectar robbers on the reproductive fitness of Jacaranda rugosa , an endemic shrub of the National Park of Catimbau, in the Caatinga of Pernambuco, Brazil. Xenogamous J . rugosa flowers continuously produced nectar during the day at a rate of 1 μl·h⁻¹. Female and male Euglossa melanotricha were the main pollinators. Early morning flower visits substantially contributed to fruit set because stigmas with open lobes were almost absent in the afternoon. Ninety-nine per cent of the flowers showed damage caused by nectar robbers. Artificial addition of sugar water prolonged the duration of flower visits of legitimate flower visitors. Removal of nectar, simulating the impact of nectar robbers, resulted in shorter flower visits of euglossine bees. While flower visits of nectar-robbing carpenter bees ( Xylocopa frontalis , X . grisescens, X . ordinaria) produced only a longitudinal slit in the corolla tube in the region of the nectar chamber, worker bees of Trigona spinipes damaged the gynoecium in 92% of the flowers. This explains the outstandingly low fruit set (1.5%) of J. rugosa in the National Park of Catimbau.     

Pollination, Pollen Viability and Pistil Receptivity in Cucurbita pepo

Cucurbita pepo carries male and female flowers on the same plant,and is pollinated by nectar-collecting bees. The nectaries aredimorphic in the two sexes and pollen is loaded and unloadedas the bees gain access to the nectar. Both types of flowerare open for only 6 h (from 0600 h to 1200 h); male flowersopen and close half an hour earlier than female flowers. Thelatter produce more nectar and are visited more often by thebees than the male flowers. Pollen viability determined by fluoresceindiacetate (fluorochromatic reaction) decreases by 20% duringanthesis and more rapidly after the flower closes. This decreaseis due to dehydration of the grain, especially around the porewhere the intine is exposed. An unusual feature of this speciesis that the grains do not dehydrate before anther dehiscence.Female receptivity has two aspects, that of the stigma lasting4 d, and that of the ovules lasting 2 d. The receptivity ofthe two sexes and the short period of anthesis are discussedin the light of the reproductive ecology of the species.Copyright1993, 1999 Academic Press Entomophilous pollination, anthesis, nectars, pollen viability, female receptivity, bees, pollinator efficiency, courgette Cucurbita pepo, Cucurbitaceae     

Native or Exotic? Double or Single? Evaluating Plants for Pollinator-friendly Gardens

In a series of dawn-to-dusk studies, we examined the natureand accessibility of nectar rewards for pollinating insectsby monitoring insect visits and the secretion rate and standingcrop of nectar in the British native plant species Salvia pratensis,Stachys palustris, S. officinalis, Lythrum salicaria, Linariavulgaris, the non-native Calendula officinalis, Petunia x hybrida,Salvia splendens, and the possibly introduced Saponaria officinalis.We also compared single with double variants ofLotus corniculatus, Saponaria officinalis, Petunia x hybrida andCalendula officinalis. All the British species studied are nectar-rich and are recommendedfor pollinator-friendly gardens. They showed maximal secretionrates of about 10–90 µg sugar per flower h^-1, andmost had mean standing crops of about 5–60 µg sugarper flower. In all British species studied, the corolla wasdeep enough for the relatively long-tongued bumblebee Bombuspascuorum, but the shallower flowers of Lythrum salicaria werealso much visited by shorter-tongued bees and hoverflies, aswell as by butterflies. The exotic Salvia splendens, presumablycoevolved with hummingbirds in the Neotropics, has such deepflowers that British bees cannot reach the nectar except bycrawling down the corolla. With a secretion rate approaching300 µg sugar per flower h^-1and little depletion by insects,S. splendens accumulated high standing crops of nectar. S. splendens,and single and double flowers of the two probably moth-pollinatedspecies Petunia x hybrida and Saponaria officinalis, receivedfew daytime visits despite abundant nectar but Calendula waswell visited by hoverflies and bees. We compared single anddouble variants of Lotus corniculatus,Petunia x hybrida andCalendula officinalis, and also Saponaria officinalis, the lastbeing probably introduced in Britain (Stace, 1997 New floraof the British Isles. 2nd edn. Cambridge: Cambridge UniversityPress). In Petunia, Saponaria and Lotus, double flowers secretedlittle or no nectar. In Calendula, where doubling involved achange in the proportion of disc and ray florets rather thanmodification of individual flower structure, double and singlecapitula had similar standing crops of nectar. Except inCalendula, exotic or double flowers were little exploited by insect visitors.In the exotics, this was probably due to the absence or scarcityof coevolved pollinators, coupled, in double flowers, with theabsence of nectar. Copyright 2001 Annals of Botany Company Salvia pratensis, Salvia splendens, Stachys palustris, Stachys officinalis,Lythrum salicaria , Linaria vulgaris, Lotus corniculatus, Saponaria officinalis,Petunia x hybrida, Calendula officinalis, wild flowers, double flowers, gardens, nectar, secretion rate, standing crop, pollinators, bumblebees, Bombus, honeybees, Apis, hoverflies, butterflies,Anthidium manicatum     

Risk‐averse inflorescence departure in hummingbirds and bumble bees: could plants benefit from variable nectar volumes?

Most hermaphroditic, many-flowered plants should suffer reduced fitness from within-plant selfing (geitonogamy). Large inflorescences are most attractive to pollinators, but also promote many flower visits during a single plant visit, which may increase selfing and decrease pollen export. A plant might avoid the negative consequences of attractiveness through modification of the floral display to promote fewer flower visits, while retaining attractiveness. This report shows that increasing only the variance in nectar volume per flower results in fewer flower visits per inflorescence by wild hummingbirds ( Selasphorus rufus ) and captive bumble bees ( Bombus flavifrons ) foraging on artificial inflorescences. Inflorescences were either constant (all flowers contained the same nectar volume) or variable (half the flowers were empty, the other half contained twice as much nectar as in the constant flowers). Both types of inflorescence were simultaneously available to foragers. Risk-averse foraging behaviour was expressed as a patch departure preference: birds and bees visited fewer flowers on variable inflorescences, and this preference was expressed when resource variability could be determined only by concurrent sampling. When variance treatments were clearly labelled with colour and offered to hummingbirds, the departure effect was maintained; however, when preference was measured by inflorescence choice, birds did not consistently prefer to visit constant inflorescences. The reduced visitation lengths on variable inflorescences by both birds and bees documented in this study imply that variance in nectar production rates within inflorescences may represent an adaptive trait to avoid the costs of geitonogamy.     

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.

     

The role of insect water balance in pollination ecology: Xylocopa and Calotropis

Summary Two carpenter bees (Xylocopa spp.) in southern Israel both use the asclepiad Calotropis procera as a primary nectar source. This plant genus is coevolved with carpenter bees, and aspects of the insect-flower interaction in Israel suggest that the smaller bee, X. sulcatipes, is the natural co-adapted pollinator, a view borne out by the geographical distributions of the species concerned. There are significant mismatches between the plant and the larger X. pubescens, involving physical fit and behaviour. These mismatches are particularly evident when the physiologies of the bees and the plant are considered. The different sizes and colours of the two bees lead to different daily activity patterns, only X. sulcatipes being thermally suited to, and thus abundant at, times of maximum nectar production by Calotropis. Similarly the water requirements of X. sulcatipes are finely balanced with the water production in the floral nectar; this bee gains just enough water when foraging to restore its blood concentration and production in the floral nectar; for deposition in the nest. X. pubescens does not incur net water loss in flight and gains too much water from Calotropis flowers, necessitating copious urination and tonguelashing. Hence physiological information can be of use in deciphering insect-plant coevolutionary patterns, and the water component of nectar is confirmed as a potentially major determinant of foraging activities. The circumstances where this will be particularly true, and cases where it may not apply, are discussed.     

Pollen in Bee-Flower Relations Some Considerations on Melittophily*

Pollen and nectar are usually lumped together as floral rewards for pollinating bees, but they play totally different roles for flowers and bees (Table 1), as well as in the relationship between them. While flowers are specialized for certain pollinators via nectar, bees specialize on certain flowers via pollen. While flowers need pollen as a prerequisite for pollination, it is the essential larval food in bees. Thus, there is a strong competition between them for pollen. Foraging for pollen must be divided into three phases: uptake in the flower, reloading into and homeward transport within a carrying container. Bees have specializations for transport but hardly any for pollen uptake - and thus for pollination. Bees actively harvesting pollen usually do not pollinate. This only happens as a consequence of contamination of the bee by pollen. From these data a scenario is provided for the evolution of bees and bee flowers. Specialized bee flowers are often characterized by their ability to hide pollen from the bees and at the same time use them as optimal pollinators. If the relationship of bees and flowers is mutualistic at all it is best described as a balanced mutual exploitation.     

Asymmetric competition for nectar between a large nectar thief and a small pollinator: an energetic point of view

There are two alternative hypotheses related to body size and competition for restricted food sources. The first one supposes that larger animals are superior competitors because of their increased feeding abilities, whereas the second one assumes superiority of smaller animals because of their lower food requirements. We examined the relationship between two unrelated species of different size, drinking technique, energy requirements and roles in plant pollination system, to reveal the features of their competitive interaction and mechanisms enabling their co-existence while utilising the same nectar source. We observed diurnal feeding behaviour of the main pollinator, the carpenter bee Xylocopa caffra and a nectar thief, the northern double-collared sunbird Cinnyris reichenowi on 19 clumps of Hypoestes aristata (Acanthaceae) in Bamenda Highlands, Cameroon. For comparative purpose, we established a simplistic model of daily energy expenditure and daily energy intake by both visitor species assuming that they spend all available daytime feeding on H. aristata. We revealed the energetic gain–expenditure balance of the studied visitor species in relation to diurnal changes in nectar quality and quantity. In general, smaller energy requirements and related ability to utilise smaller resources made the main pollinator X. caffra competitively superior to the larger nectar thief C. reichenowi. Nevertheless, sunbirds are endowed with several mechanisms to reduce asymmetry in exploitative competition, such as the use of nectar resources in times of the day when rivals are inactive, aggressive attacks on carpenter bees while defending the nectar plants, and higher speed of nectar consumption.     

Are nectar-robbers mutualists or antagonists?

As exploiters of plant-pollinator mutualisms, nectar-robbers remove rewards (nectar) without providing pollination services. Though one might expect nectar-robbing to be costly to plants, it may instead benefit plants by indirectly increasing pollen dispersal. I investigated the direct effects of nectar-robbing bees (Xylocopa californica) on floral rewards and behaviors of pollinators visiting desert willow (Chilopsis linearis) and indirect effects of robbing on the reproductive success of the plant. Nectar-robbers reduced nectar; while unrobbed and robbed flowers were equally likely to contain nectar, nectar volumes were smaller in robbed flowers with nectar. Apis mellifera (honeybees), ineffective pollinators in terms of pollen deposition, avoided robbed flowers. In contrast, Bombus sonorus (bumblebees), effective pollinators, did not avoid robbed flowers. While bumblebees tended to spend less time in robbed flowers, the time that they spent in flowers was not correlated with pollen deposition. Using powder mimicking pollen, I found that on some days, powder was dispersed farther or to more flowers from robbed flowers, indicating that robbing may sometimes benefit plants by increasing male reproductive success. Powder movement suggested that the effect of robbing on male reproductive success ranged from costly to beneficial. The outcome for flowers that were marked early each morning was a function of prevalence of robbing and abundances of effective pollinators, but not a function of spatial variability among trees in prevalence of robbing or the abundance of ineffective honeybees. Unlike powder dispersal, female reproductive success, measured by fruit set and the number of pollen tubes growing in styles, was not affected by robbing. Thus, robbers did not reduce plants female reproductive success either directly by damaging flowers or indirectly by reducing pollen deposition by pollinators. Overall, this study indicates that nectar-robbers were not often costly to plants, and sometimes even benefited plants.     

Does the Flower Constancy of Bumble Bees Reflect Foraging Economics?

We examined the effects of floral reward level and spatial arrangement on the propensity of bumble bees to exhibit flower constancy. In three separate experiments, we compared the flower constancy of bees on dimorphic arrays of blue and yellow flowers that differed either in reward concentration, reward volume, or inter‐flower distance. Overall, flower choice patterns varied among bees, ranging from random selection to complete constancy. When flowers contained greater reward volumes and were spaced farther apart, bees showed less flower constancy and more moves to closely neighbouring flowers. Changes in reward concentration had no effect on flower constancy; however, more dilute rewards produced shorter flight times between flowers. In addition, there was a strong positive relationship between degree of flower constancy and net rate of energy gain when flowers were spaced farther apart, indicating that constant bees were more economic foragers than inconstant bees. Together, these results support the view that the flower constancy of pollinators reflects an economic foraging decision.     

Assessment of pollen reward and pollen availability in Solanum stramoniifolium and Solanum paniculatum for buzz‐pollinating carpenter bees

The two widespread tropical Solanum species S. paniculatum and S. stramoniifolium are highly dependent on the visits of large bees that pollinate the flowers while buzzing them. Both Solanum species do not offer nectar reward; the rewarding of bees is thus solely dependent on the availability of pollen. Flower visitors are unable to visually assess the amount of pollen, because the pollen is hidden in poricidal anthers. In this study we ask whether and how the amount of pollen determines the attractiveness of flowers for bees. The number of pollen grains in anthers of S. stramoniifolium was seven times higher than in S. paniculatum. By contrast, the handling time per five flowers for carpenter bees visiting S. paniculatum was 3.5 times shorter than of those visiting S. stramoniifolium. As a result foraging carpenter bees collected a similar number of pollen grains per unit time on flowers of both species. Experimental manipulation of pollen availability by gluing the anther pores showed that the carpenter bees were unable to detect the availability of pollen by means of chemical cues before landing and without buzzing. Our study shows that the efficiency of pollen collecting on S. paniculatum is based on large inflorescences with short between‐flower search times and short handling time of individual flowers, whereas that of S. stramoniifolium relies on a large amount of pollen per flower. Interestingly, large carpenter bees are able to adjust their foraging behaviour to drastically different strategies of pollen reward in otherwise very similar plant species.     

Nectarivore foraging ecology: rewards differing in sugar types

Abstract.

• 1 Honey bees, visiting artificial flower patches, were used as a model system to study the effects of sugar type (sucrose, glucose, fructose, and mixed monosaccharide), caloric reward, and floral colour on nectarivore foraging behaviour. Observed behaviour was compared to the predictions of various (sometimes contradictory) foraging models.
• 2 Bees drank indiscriminately from flowers in patches with a blue-white flower dimorphism when caloric values of rewards were equal (e.g. 1M sucrose in both colours; 1 M sucrose versus 2 M monosaccharide of either type), but when nectar caloric rewards were unequal, they switched to the flower colour with the calorically greater reward.
• 3 In yellow-blue dimorphic flower patches, on the other hand, bees did not maximize caloric reward. Rather, bees were individually constant, some to blue, others to yellow, regardless of the sugar types or energy content of the rewards provided in the two flower morphs.
• 4 The results suggest that optimal foraging theory (maximization of net caloric gain per unit time) is a robust predictor of behaviour with regard to the sugar types common to nectars; such optimal foraging is, however, limited by a superstructure of individual constancy.

     

Relationship between floral tube length and nectar robbing in Duranta erecta L. (Verbenaceae)

Although nectar robbing is a common phenomenon in plant species with tubular flowers or flowers with nectar spurs, the potential effect of this illegitimate interaction on plant reproductive success has not received the deserved attention. In the present study, we analysed the functional relationship between flower morphology and nectar robbing, and examined the reproductive consequences of the interaction in a population of Duranta erecta (Verbenaceae) on the island of Cuba. The results show that nectar robbing is conducted by the carpenter bees Xylocopa cubaecola and affects up to 44% of flowers in the studied population. However, not all the flowers have the same probability of being robbed. The chance of flowers being robbed increases with flower length and flower diameter. Moreover, nectar robbing significantly decreases the chance that flowers will set fruit. Also, the impact of nectar robbing on the probability of flowers to set fruits is dependent on the plant. We suggest that nectar robbing may represent an opposite selective force that balances the selection for longer corollas often imposed by pollinators specializing in visiting tubular flowers. Such a relationship with nectar robbers would have obvious implications for the evolution of tubular or closed flowers. This preliminary finding deserves further research in light of the ecological and evolutionary consequences of nectar robbing in tubular flowers.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 392–398.