THE POLLINATION ECOLOGY OF PEDICULARIS ON MOUNT RAINIER

Six native species of Pedicularis in Mount Rainier National Park in Washington State were studied for their reproductive relationships with animal pollinators. Cinematographic and stereophotographic records revealed pollination of the nectariferous P. bracteosa and P. rainierensis by upright, nectar-foraging queens and workers of five bumblebee (Bombus Latr.) species and by inverted workers scraping pollen from anthers concealed within the corolla galea. The nectarless, rostrate flowers of P. contorta, P. groenlandica, P. ornithorhyncha, and P. racemosa were pollinated by pollen-foraging workers and occasional queens virbrating pollen from concealed anthers. Insect exclosure methods revealed complete absence of fruiting in the absence of insects, and pollinator collections further indicate obligate dependence of the plants upon bumblebees for their sexual reproduction. Analysis of corbicular pollen loads from pollinators suggested that pollinator species are not monolectic but that individual pollinators range from monolectic to polylectic. Measurements indicated limited correlations between lengths of corolla tubes and tongues of nectar-foraging insects. Each nectarless Pedicularis species occupied a different, specific habitat, but P. bracteosa and the endemic P. rainierensis were sympatric in part. Each species had a unique spectral reflectance pattern from the corolla. Proximity of habitats and overlap of blooming periods of all Pedicularis species eliminate the possibility of contemporary geographic or phenological reproductive isolation. It is suggested that behavioral interactions of the plants and their insect pollinators may have been instrumental in the past in reproductively isolating these species, hybrids of which are unknown.     

Deceptive pollination of the Lady's Slipper Cypripedium tibeticum (Orchidaceae)

To test whether the nectarless flowers of Cypripedium tibeticum attract pollinators through mimicry like the allied species C. macranthos var. rebunense, pollination biology of C. tibeticum was investigated in western China. Although C. tibeticum was also pollinated by bumble bee queens, i.e. Bombus lepidus , B. lucorum and B. hypnorum , no special, rewarding model plants were found in the habitat. Field experimentation confirmed that the flowers were self-compatible but insects were required to transfer orchid pollen to the stigma. Both Bombus queens and workers were visitors, but queens were much more frequent than workers and only queens were effective pollinators. Floral functional morphology analysis showed that it was large queens rather than small workers that fitted well with the flowers of C. tibeticum. With the faint sweet-fruity scent, the minor floral fragrance compound, ethyl acetate, probably plays a role in attracting bumble bees by food deception. The dark flowers with the inflated, trap-like labellum are hypothesized to mimic the nest site of queens. Therefore, bumble bee queens tend to be attracted by C. tibeticum through nest site mimic combined with food deception. Considering that the co-blooming flowers of C. flavum are pollinated by the Bombus workers, and C. smithii pollinated by a queen, we suggest that using the same bumblebees with different body sizes as the pollinators is the main reproductive isolation between interfertile C. tibeticum and C. flavum, while C. tibeticum and C. smithii tend to hybridize naturally.     

POLLEN-FORAGING BEHAVIOR OF BOMBUS IN RELATION TO POLLINATION OF NOTOTRIBIC FLOWERS

Cinematographic records of the pollen-foraging behavior of six Bombus species on Linaria vulgaris, Lobelia siphilitica, and Antirrhinum majus revealed a generally uniform pattern of pollen deposition by the nototribic flowers on the frons and scutum of the foragers. A forward sweeping motion of the tarsal brushes of their middle legs removed some of this pollen which was transferred to the tibial corbicula of the hind legs. Some insects foraged in an inverted position. An analysis of 804 pollen samples from frons, scutum, and corbiculum of 331 from a total of 395 Bombus visitors, representing eight of the nine native species, on the three plants indicated a high proportion of proper pollen being collected in addition to 15 kinds of foreign pollen. The pollen-sweeping behavior of Bombus on nototribic flowers is suggested as an important means of pollen-foraging apart from its utility in insect grooming. Pollen-foraging insects are considered important pollinators of these nototribic flowers.     

The most effective pollinator revisited: pollen dynamics in a spring-flowering herb

Most flowers are visited by a wide range of potential pollinators. However, their efficiency in pollen removal and deposition, and other behavioural factors affecting pollination effectiveness may greatly differ among taxa, and even individuals. Fritillary (Fritillaria meleagris L., Liliaceae) is a spring-flowering, critically endangered plant in the Polish flora, red-listed in most of the European countries of its range. Based on indirect evidence, that is, body pollen loads, visitation frequency and seasonal abundance, it is estimated that its key pollinators are queen bumblebees, but, as shown in the literature, the largest Fritillaria pollen loads are carried by solitary bees. To study pollinator effectiveness for floral visitors to F. meleagris, we performed a garden experiment, where we analysed pollen deposition and assessed pollen removal per single flower-visit in the plant. Similarly to field conditions reported in the literature, our experimental plants were serviced by nectar-seeking bumblebee queens and two taxa of solitary bees, small pollen-collecting Andrena and large, nectar-seeking Anthophora males. When “quality” component was addressed, despite the character of visits, insects from all groups deposited more pollen than was found on unvisited flowers, but they did not differ significantly from each other in pollen deposition on virgin stigmas. We also found some differences in pollen removal both within- and among-visitor species and control flowers, unfortunately due to extremely high variation of the results they were all statistically insignificant. However, when “quantity” component of insect performance was concerned, we observed that over 81 % of visits were by bumblebees. Bombus queens stayed on flowers significantly less time than small Andrena individuals (13 % of recorded visits) and equally long as Anthophora males (only 6 % of visits). We conclude that although all the visitor groups can pollinate the flowers of F. meleagris, bumblebee queens indeed proved to be the most effective pollinators of the plant, when both quality and quantity components of pollination are concerned.     

Pollen-Foraging Dynamics of Subalpine Bumblebees (Bombus Latr.)

Abstract A 23-year study on Mount Rainier of nectar-foraging hummingbirds and pollen-foraging bumblebees (Bom-bus Latr.) indicated a constant relative frequency of Bombus species in the area and a wide occurrence of insect species on flowers plus a wide range of flower species pollinated by a single insect. Early-blooming plants had a high queen/worker pollinator ratio, while workers predominated on later-blooming plants. No altitudinal difference in distribution of Bombus species occurred. Analysis of 955 corbicular pollen loads from 1158 bumblebees collected yielded 49.5% monolecty for queens and 34.5% for workers with a wider range for queens than for workers. Colors of corollas photographed in visible and ultraviolet light corresponded to the visual spectra of animal pollinators. In the stable plant-pollinator community investigated, pollinating insect activity appears phenologically controlled by floral anthesis and behaviorally related to floral form and function. Bumblebee tongue length, however. Is not a valid determinant for foraging niche separation except in Castilleja with a deep corolla tube accommodating long tongues of hummingbirds and bumblebees. It is concluded that in a biotic community well-coordinated with the physical environment, limited flower constancy of pollen-foraging pollinators appears to contribute a necessary degree of adaptive versatility through sharing of insect and floral resources. Flower constancy is primarily a function of the circumstance in which a pollinator operates and not simply a characteristic of the pollinator itself.     

THE POLLINATION DYNAMICS OF SYMPATRIC SPECIES OF PEDICULARIS (SCROPHULARIACEAE)

Sympatric populations of Pedicularis oederi, P. cystopteridifolia, and P. groenlandica on the Beartooth Plateau (Montana) were obligately dependent on Bombus pollinators. Their corolla colors were mutually distinct to insect vision, but their nectars had identical sugar components. Analysis of corbicular pollen loads of pollinating insects indicated a high degree of polylecty. Queen/worker ratios of pollinators on plant species corresponded to the parallel phenological sequences of plant blooming and insect caste development. The total number of individual pollinators of each Bombus species on all plants was directly related to the number of plant species it pollinated, but a comparable relationship between the number of pollinators on a plant species to the number of Bombus species pollinating it was not found. Morphological and behavioral correspondence of floral mechanisms and pollinators form an integral part of the general pattern of coadaptive evolution of the floral ecology of Pedicularis in North America known from previous studies. Reproductive isolation between Pedicularis species in this study is attributed to internal barriers. Wide overlaps of blooming seasons, proboscis lengths of bumblebee species and castes, and pollinator species on Pedicularis species suggest resource sharing, rather than resource partitioning or competition for resources among plant and insect species and individuals.     

Pollination ecology in sympatric winter flowering Helleborus (Ranunculaceae)

We studied biotic and abiotic factors that influence pollination in two sympatric winter flowering species. Helleborus foetidus and Helleborus bocconei flower simultaneously in winter. Although climatic conditions are not favorable for biotic pollination both species rely mainly on large bees of the genus Bombus. At the beginning of flowering, harsh climatic conditions are restrictive for insect visits. As flowering continues and temperatures rise, pollinator activity increases. The two plant species share pollinators that visit them indiscriminately. The flowers of the two species differ in form and insects visit H. foetidus by inserting their heads and H. bocconei ventrally: pollen load on insects is highly specialized. With the arrival of spring, many other species start to bloom and in spite of the large number of flowers still on the plants insects abandon Helleborus species. At the end of spring increasing biotic interactions take away pollinators from the Helleborus species.     

Pollination of the lady's slipper Cypripedium henryi Rolfe (Orchidaceae)

The pollination ecology of Cypripedium henryi Rolfe, a slipper orchid endemic to west China, was investigated, and its floral shape, size, colour, and scent were analysed. Examination of the breeding system suggests that the flowers are self-compatible, but need pollen vectors for successful reproduction. The flower is rewardless; over 15 insects belonging to Araneida, Hymenoptera, Diptera, Lepidoptera, and Coleoptera were recorded as flower visitors, but most only alighted or rested on the flower. In the total 32 h of observations over 2 years, female Lasioglossum bees were found to be the most frequent visitors and the only pollinators. They showed a high visitation frequency and, surprisingly, re-visited the same flowers frequently. Cypripedium henryi probably attracts pollinators visiting the flowers through general food deception (odour components, colour, false nectar guides), as well as special structures (slippery labellum, slippery staminode). Although three Lasioglossum species visited the flowers, only L. sauterum Fan et Ebmer was found with pollen. Lasioglossum flavohirtum Ebmer was large and climbed out from the entrance. Morphologically, L. sichuanense Fan et Ebmer could be considered as a potential pollinator, but the collected specimens were found to have no pollen of C. henryi on their bodies. It was speculated that the particular floral scent of C. henryi discouraged the entrance of L. sichuanense bees. Lasioglossum sauterum was matched morphologically to the flower, but not all of the visitations resulted in effective pollinations, as some flowers of C. henryi were frequently re-visited and the pollen mass had been taken away by bees on previous visitations. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 491–499.     

POLLEN AND GENE DISPERSAL: THE INFLUENCES OF COMPETITION FOR POLLINATION

Pollinators that forage indiscriminately can transfer pollen from one species to another, reducing the amount that reaches conspecific flowers. I present evidence that the presence of another plant species visited by the same pollinators can also reduce pollen dispersal distances and outcrossing. This has the potential to influence gene flow and reproductive success. Pollen carryover and movement patterns were measured for the shared insect pollinators of Stellaria pubera and Claytonia virginica in North Carolina. Bee flies deposited similar amounts of Stellaria pollen on a series of pistillate Claytonia flowers as on a series of pistillate Stellaria flowers. In arrays of potted plants, flies and solitary bees visited most flowers on a plant before leaving and then flew to a nearby plant chosen independently of species; 95% of moves were to one of 12 nearest neighbors. These measures of pollen carryover and movement patterns were used in a set of computer simulations to predict pollen dispersal distances. The simulations suggested that C. virginica substantially reduces outcrossing and pollen flow in S. pubera. These predictions were tested by tracking dye movement from anthers in populations of potted plants. Addition of C. virginica reduced the mean squared distance moved by dye to receptive S. pubera flowers by 23% and reduced the amount of dye moved by 51%. The estimated pollen component of gene flow was also much lower in a natural population of 5. pubera mixed with C. virginica than in the synthetic single-species populations.     

Effectiveness of short‐tongued bees as pollinators of apparently ornithophilous New Zealand mistletoes

Abstract The flowers of two species of threatened New Zealand mistletoes (Peraxilla tetrapetala and Peraxilla colensoi, Loranthaceae) have explosive buds that do not open unless force is applied by birds or two species of native short‐tongued bees. Opened flowers are visited by a variety of birds and insects. Although both species of Peraxilla conform to a pollination syndrome of ornithophily, bees may be effective alternative pollinators. We investigated the effectiveness of bees and birds as pollinators of P. colensoi at one site and P. tetrapetala at two sites in the South Island. Bees and other insects outnumbered birds as flower visitors at all three sites. By excluding birds with wire cages, we showed that two bee species regularly open flowers of P. tetrapetala, but only rarely open flowers of P. colensoi. Few pollen grains were deposited when either birds or bees opened buds, so opening buds was not by itself sufficient for adequate pollination. Instead, pollen continued to accumulate over the next 6 or 7 days, even inside cages that excluded birds. Both populations of P. tetrapetala were regularly pollen‐limited, but in different ways. At Ohau, opened flowers gained enough pollen to produce seeds, but many buds were not opened and hence failed to set seed. In contrast, at Craigieburn, nearly all buds were opened, but many of these did not receive enough pollen. These results demonstrate that native bees can partially replace birds as pollinators of mistletoes, despite their apparent ornithophilous syndrome. Ongoing reductions in New Zealand forest bird numbers means that the service bees provide may be important for the long‐term future of these plants.     

Three wild Lithuanian strawberry species and their pollinators

The European wild strawberry species Fragaria vesca L., Fragaria viridis Weston, and Fragaria moschata Weston are sympatric and bloom concurrently, introducing the possibilities of cooperation and competition for pollinators. We studied the pollination biology of strawberries that were grown in rectangular plots at a field site near Vilnius city in Lithuania. The number of open flowers, the number of insects visiting flowers, and the plot area covered by strawberry plants were recorded. Wild strawberries were visited by solitary bees (38.5%), flies (31.4%), and other hymenopterans (21%; ants accounted for approximately 98% of this group). Coleopterans, lepidopterans and honey bees together represented less than 10% of the insects recorded on strawberries. Fragaria viridis and F. moschata flowers were more attractive to pollinators than those of F. vesca. The main visitors of F. viridis were small solitary bees; those of F. moschata were ants, while F. vesca flowers were visited by solitary bees, ants and flies in comparable numbers. We discuss competition for pollinators and the possible impact of climate fluctuation on strawberry-pollinator interactions.     

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.

     

Mechanisms that limit pollinator range in Ericaceae

Studies were conducted in 2001-2003 at Valdai National Park (Novgorod region) and at the Zvenigorod biological station of Moscow State University. The morphology of flowers, flowering dynamics and composition of insect visiting flowers of Ericaceae species: Andromeda polifolia, Chamaedaphne calyculata, Ledum palustre, Oxycoccus palustris, Vaccinium myrtillus, V. uliginosum, and V. vitis-idaea L. were studied. Some species of insects visiting flowers were excluded from the list of pollinators on the basis of observation on their behavior. L. palustre was visited mainly by flies where as other investigated species were visited mainly by bumblebees. In some cases bumblebees were the only visitors of the investigated plants. Mechanisms that protect flowers from flies and short-tongued solitary bees visits and ensure a best pollination by bumblebees are various among different species of Ericaceae. Efficiency of nectary protection also differs among different plant species and is defined by particularities of their habitats and flowering phenology. As far as all species of this family during the flowering are dominants in typical habitats, a competition for the pollination with species of other families in most cases is megligible. Flowering periods of V. vitis-idaea and V. myrtillus in forest ecosystems overlapped weakly. Moreover, V. myrtillus is pollinated mainly by bumblebee queens where as pollinators of V. vitis-idaea are bumblebee workers, solitary bees and horse flies. The other investigated plant species inhabit only oligotrophic peat bogs. Thery are pollinated by bumblebees but periods of flowering are not overlapped and consequently follow one after another. L. palustre and V. uliginosum flower simultaneosly but they are pollinated by different pollinators.     

Bees assess pollen returns while sonicating Solanum flowers

Summary Can bees accurately gauge accumulating bodily pollen as they harvest pollen from flowers? Several recent reports conclude that bees fail to assess pollen harvest rates when foraging for nectar and pollen. A native nightshade (Solanum elaeagnifolium Cavanilles) that is visited exclusively for pollen by both solitary and social bees (eg. Ptiloglossa and Bombus) was studied in SE Arizona and SW New Mexico. The flowers have no nectaries. Two experiments were deployed that eliminated pollen feedback to the bees by experimentally manipulating flowers prior to bee visits. The two methods were 1) plugging poricidal anthers with glue and 2) emptying anthers of pollen by vibration prior to bee visitation. Both experiments demonstrated that bees directly assess pollen harvest on a flower-by-flower basis, and significantly tailor their handling times, number of vibratile buzzes per flower and grooming bouts according to the ongoing harvest on a given flower. In comparison to experimental flowers, floral handling times were extended for both Bombus and Ptiloglossa on virgin flowers. Greater numbers of intrafloral buzzes and numbers of times bees groomed pollen and packed it into their scopae while still on the flower were also more frequent at virgin versus experimental flowers. Flowers with glued andreocia received uniformly brief visits from Bombus and Ptiloglossa with fewer sonications and virtually no bouts of grooming. Curtailed handling with few buzzes and grooms also characterized visits to our manually harvested flowers wherein pollen was artificially depleted. Sonicating bees respond positively to pollen-feedback while harvesting from individual flowers, and therefore we expect them to adjust their harvesting tempo according to the currency of available pollen (standing crop) within Solanum floral patches.     

Inflorescence Height Affects Visitation Behavior of Bees–A Case Study of an Aquatic Plant Community in Bolivia1

We studied the bee fauna visiting a plant community of 10 species of flowering aquatic plants in an inundated savanna region in Bolivia. In total we observed 36 bee species in 17 genera at the flowers. Cluster analysis of the similarities among the plant species in terms of their visitor spectra showed a division into two groups: plants with inflorescence heights shorter than the grass height and plants with inflorescences projecting out of the surrounding vegetation. Larger bees of the genera Apis, Melipona, Bombus, and Xylocopa were observed only at flowers above the surrounding vegetation. Smaller, mainly solitary bees (e.g., Augochlorella, Ancyloscelis) visited flowers in the dense vegetation near the water surface. Analyses of the pollen loads revealed that most individuals were highly flower constant. When bees carried different pollen types, it was generally pollen from flowers within a single stratum. We discuss specialization, flower constancy, competition, and different foraging strategies as possible reasons for stratum fidelity.     

The endangered Iris atropurpurea (Iridaceae) in Israel: honey-bees,night-sheltering male bees and female solitary bees as pollinators

Background and Aims

The coastal plain of Israel hosts the last few remaining populations of the endemic Iris atropurpurea (Iridaceae), a Red List species of high conservation priority. The flowers offer no nectar reward. Here the role of night-sheltering male solitary bees, honey-bees and female solitary bees as pollinators of I. atropurpurea is documented.

Methods

Breeding system, floral longevity, stigma receptivity, visitation rates, pollen loads, pollen deposition and removal and fruit- and seed-set were investigated.

Key Results

The main wild pollinators of this plant are male eucerine bees, and to a lesser extent, but with the potential to transfer pollen, female solitary bees. Honey-bees were found to be frequent diurnal visitors; they removed large quantities of pollen and were as effective as male sheltering bees at pollinating this species. The low density of pollen carried by male solitary bees was attributed to grooming activities, pollen displacement when bees aggregated together in flowers and pollen depletion by honey-bees. In the population free of honey-bee hives, male bees carried significantly more pollen grains on their bodies. Results from pollen analysis and pollen deposited on stigmas suggest that inadequate pollination may be an important factor limiting fruit-set. In the presence of honey-bees, eucerine bees were low removal–low deposition pollinators, whereas honey-bees were high removal–low deposition pollinators, because they removed large amounts into corbiculae and deposited relatively little onto receptive stigmas.

Conclusions

Even though overall, both bee taxa were equally effective pollinators, we suggest that honey-bees have the potential to reduce the amount of pollen available for plant reproduction, and to reduce the amount of resources available to solitary bee communities. The results of this study have potential implications for the conservation of this highly endangered plant species if hives are permitted inside reserves, where the bulk of Oncocyclus iris species are protected.     

THE POLLINATION ECOLOGY OF ASTRAGALUS CIBARIUS AND ASTRAGALUS UTAHENSIS (LEGUMINOSAE)

Astragalus cibarius and A. utahensis are common perennial species of a widespread legume genus. The pollination of Astragalus has been briefly discussed in the literature, but little work has been done on species in the intermountain West. This study was conducted from 1970–1973 in Utah with mixed and single species populations. The flowers of both species were homogamous and papilionaceous, but the species were different as to color, size, and ultraviolet reflectance. Astragalus cibarius usually flowered 10 days ahead of A. utahensis, but both species flowered earlier than most other plants in the community. Bagging experiments indicated both species were strongly allogamous. Exclosure studies indicated both species relied on insects as pollen vectors. Of the 44 insect species which were observed visiting flowers, only 14 carried Astragalus pollen, and the pollinator fauna varied between study sites. Pollen quantities and distributions on Diptera and Coleoptera indicated a poor potential for pollination. Floral structure, pollen distribution and quantity, and behavior implied that large bees of the families Apidae and Anthophoridae were the primary pollinators. These bees visited only one species of Astragalus when the plants occurred in mixed populations; this constancy may have been related to relative flower abundance. Non-pollinating floral foragers affect other phases of Astragalus life history.     

Foraging Behaviour of Wild Bees at Hot Pepper Flowers (Capsicum annuum) and its Possible Influence on Cross Pollination

Native bees are effective pollinators of hot pepper plants,Capsicum annuum. In four gardens in south-central Brazil (Patosde Minas and Brasília) flowers of three cultivars receivedvisits from 16 species of bees in eight genera: Hylaeus(Colletidae),Dialictus, Halictus, Augochlora,Augochloropsis and Ceratalictus(Halictidae),Exomalopsis andBombus (Apidae). No other insects were observedto visit the flowers. Some species of bee occurred in more thanone garden. Individual bees gathered a full pollen load from18 to 47 flowers and visited one to eight plants on a singleforaging trip. In 76 shifts between plants, the bees made nineswitches between cultivars. It is suggested that small nativespecies of bees pollinate the flowers effectively and that theirsmall foraging areas are important in keeping the cultivarsof both hot and sweet peppers genetically distinct where severalcultivars are grown close together. Copyright 2000 Annals ofBotany Company Capsicum annuum, hot pepper, native bees, fruit set, Brazil     

FORAGING BEHAVIOR OF BOMBUS (HYMENOPTERA: APIDAE) IN RELATION TO AQUILEGIA POLLINATION

Observations on native and cultivated plants indicated that the ruby-throated hummingbird (Archilochus colubris L.) and at least four species of Bombus are probably effective pollinators of Aquilegia canadensis L. in southeastern Wisconsin. Bombus affinis Cresson perforated more than 90% of 2173 nectar spurs of flowers at full anthesis on wild plants. Cinematographic and stereo-photographic records of insects revealed a general behavioral pattern of mandibular perforation following exploration by antennae, maxillae, and tongue. Pollen-foraging behavior on 19 cultivated Aquilegia species and varieties was uniform, but nectar-foraging behavior varied with flower form and position and not with color variation within one flower form. Observations on marked insects showed that workers tended to forage in one manner on one plant variety, while queens foraging in one or more ways visited up to five different flower forms on one flight. Initial observations on neonate workers from a cultivated B. affinis colony suggested that a probably innate mandibular perforation behavior is not instinctively directed toward nectar-bearing floral parts, and that foraging behavior is capable of rapid establishment and considerable versatility.     

Solitary and social bees as pollinators of Wahlenbergia (Campanulaceae): single-visit effectiveness, overnight sheltering and responses to flower colour

Solitary bees often form specialised mutualisms with particular plant species, while honeybees are considered to be relatively opportunistic foragers. Thus, it may be expected that solitary bees are more effective pollinators than honeybees when foraging on the same floral resource. To test this, we studied two Wahlenbergia species (Campanulaceae) in South Africa that are visited by both social honeybees and solitary bees, and which are shown here to be genetically self-incompatible and thus reliant on pollinator visits for seed production. Contrary to expectation, the solitary bee Lipotriches sp. (Halictidae) and social bee Apis mellifera (Apidae), which were the two most frequent visitors to flowers of the study species, were equally effective pollinators in terms of the consequences of single visits for fruit and seed set. Both bee species preferentially visited female phase flowers, which contain more nectar than male phase flowers. Male solitary bees of several genera frequently shelter overnight in flowers of both Wahlenbergia species, but temporal exclusion experiments showed that this behaviour makes little contribution to either seed production or pollen dispersal (estimated using a dye particle analogue). Manipulation of flower colour using a sunscreen that removed UV reflectance strongly reduced visits by both bee groups, while neither group responded to Wahlenbergia floral odour cues in choice tests. This study indicates that while flowers of Wahlenbergia cuspidata and W. krebsii are pollinated exclusively by bees, they are not under strong selection to specialise for pollination by any particular group of bees.