The pollination efficiency of a pollinator depends on its foraging strategy,flowering phenology,and the flower characteristics of a plant species

Honey bees and stingless bees are generalist visitors of several wild and cultivated plants. They forage with a high degree of floral fidelity and thereby help in the pollination services of those plants. We hypothesized that pollination efficiency might be influenced by flowering phenology, floral characteristics, and resource collection modes of the worker bees. In this paper, we surveyed the foraging strategies of honey bees (Apis cerana, Apis dorsata, and Apis florea) and stingless bees (Tetragonula iridipennis) concerning their pollination efficiencies. Bees showed different resource gathering strategies, including legitimate (helping in pollination as mixed foragers and specialized foragers) and illegitimate (serving as nectar robbers and pollen thieves) types of flower visitation patterns. Foraging strategies are influenced by the shape of flowers, the timing of the visitation, floral richness, and bee species. Honey bees and stingless bees mainly acted as legitimate visitors in most plants studied. Sometimes honey bees served as nectar robbers in tubular flowers and stingless bees as pollen thieves in large-sized flowers. Among the legitimate categories, mixed foragers have a comparatively lower flower visitation rate than the specialized nectar and pollen foragers. However, mixed foragers have greater abundance and higher values of the single-visit pollination efficiency index (PEi) than nectar and pollen foragers. The value of the combined parameter ‘importance in pollination (PI)’ was thus higher in mixed foragers than in nectar and pollen foragers.     

Partitioning of pollen resources by two stingless bee species in the north Bahia,Brazil

Analysing the pollen stored by stingless bees allows identification of the diversity of flowering plants visited by these bees while collecting resources during a given period. However, few studies have focused on investigating the sharing of resources between species of native bees that coexist in transitional plant formations, which support conservation efforts and increase colony productivity. This study aims to describe the partitioning of pollen resources among colonies of two species of stingless bees in the state of the Bahia (Brazil) through the analysis of stored pollen. The study was conducted in neighbouring vegetation zones of the relevant species Melipona quadrifasciata anthidioides and Melipona scutellaris by collecting monthly pollen samples stored by three colonies of each species over one year. The collected samples were treated using the acetolysis technique, and the partitioning of pollen resources between the two species was investigated through quali-quantitative analysis of the treated pollen. The results show that among the 16 pollen types belonging to a particular species or group identified in the Melipona quadrifasciata anthidioides samples and the 18 identified in the Melipona scutellaris samples, 13 were shared by both species. The two most common pollen types stored by both species were from Mimosa arenosa (Fabaceae, Mimosoidae) and Eucalyptus sp. (Myrtaceae). These results demonstrate the niche similarity of pollen pasture between these two species and suggest that conservation efforts for these bee species should focus on the plant families most visited by bees.     

RELATIVE POLLINATOR EFFECTIVENESS AND EVOLUTION OF FLORAL TRAITS IN SPATHIPHYLLUM FRIEDRICHSTHALII (ARACEAE)

We studied the relative effectiveness of different pollinators of Spathiphyllum friedrichsthalii Schott for 15 months on Barro Colorado Island, Panama. Pollen-foraging stingless bees (Apidae: Trigona) made 87% of floral visits. Experiments showed that these bees pollinate flowers, and correlations of fruit- and seed-set with visitation frequencies and floral contact times suggested that they were responsible for the majority of seeds produced. Fifteen species of fragrance-foraging, male euglossine bees (Apidae: Euglossini) collectively accounted for a small portion of seed-set in fewer than 27% of the inflorescences. Neocorynura (Halictidae) were pollen thieves and were unimportant as pollinators. We propose that euglossine and stingless bees differentially influence outcrossing rates and the evolution of floral traits of S. friedrichsthalii. Foraging behavior of male euglossines should allow for more long-distance pollen flow whereas stingless bees are likely to promote near-neighbor and geitonogamous pollinations. We discuss why the prolonged male phase of anthesis in this protogynous species may be maintained through pollination by stingless bees rather than male euglossines. Furthermore, although the floral fragrance is attractive to many species of male euglossines, it attracts few individuals. This condition may represent an intermediate step in the evolution of predominant pollination by male euglossines.     

Aggressive foraging of social bees as a mechanism of floral resource partitioning in an Asian tropical rainforest

Interference competition by aggressive foraging often explains resource partitioning, but mechanisms contributing to partitioning have rarely been studied in Asian social bee guilds. Foraging of social bees at canopy flowers of Santiria laevigata (Burseraceae) and honey-water feeders was studied in a lowland mixed-dipterocarp forest in Sarawak, Malaysia. Four stingless bee species (Apidae, Meliponinae), Trigona canifrons, T.␣fimbriata, T. apicalis and T. melina, aggressively defended flower patches and feeders. At the flowers, T.␣canifrons excluded other bees only in the morning when nectar flow peaked. At the feeders, the aggression resulted in asymmetric interference competition, which produced a dominance hierarchy among seven social bee species. Interspecific partitioning of the feeders was detected in time and height but not quality. Only time of the first arrival after feeder presentation was negatively correlated with the dominance hierarchy: more aggressive species arrived at the feeders later than less aggressive species. This result suggests that a trade-off between searching ability and defensive ability at flower patches gives rise to resource partitioning in the social bee guild. Received: 22 June 1996 / Accepted: 22 November 1996     

Smelling like resin: terpenoids account for species-specific cuticular profiles in Southeast-Asian stingless bees

Insects may be unique in having a cuticle with a species-specific chemical profile. In social insects, colony survival depends not only on species-specific but also on colony-specific cuticular compounds with hydrocarbons playing an important role in the communication systems of ants, termites, wasps and bees. We investigated inter- and intraspecific differences in the composition of compounds found on the body surface of seven paleotropical stingless bee species (Apidae: Meliponini) at two different sites in Borneo (Sabah, Malaysia). Besides hydrocarbons, the body surface of all seven stingless bee species comprised terpenoid compounds, a substance class that has not been reported for chemical profiles of any social insect so far. Moreover, the chemical profile of some species differed fundamentally in the composition of terpenoids with one group (e.g. sesquiterpenes) being present in one species, but missing in another. Chemical profiles of different colonies from the same species showed the same hydrocarbon- and terpenoid compounds over different regions, as tested for Tetragonilla collina and Tetragonula melanocephala. However, chemical profiles differed quantitatively between the different colonies especially in T. melanocephala. It is likely that the terpenoids are derived from plant resins because stingless bees are known to collect and use large amounts of resins for nest construction and defence, suggesting an environmental origin of the terpenoids in the chemical profile of paleotropical stingless bees.     

Pollen use by Megalopta sweat bees in relation to resource availability in a tropical forest

1. Spatial and temporal availability of pollen helps shape bee foraging behaviour and productivity, which has been studied in great detail at the landscape level, but never in a diverse tropical forest. 2. To study the effect of spatio‐temporal variation in resource distribution on pollen use and productivity, we identified pollen from spatially explicit nest collections of two generalist sweat bees, Megalopta genalis Meade‐Waldo and M. centralis Friese, from Barro Colorado Island, Panama, a 50‐ha forest dynamics plot during the 2007 dry and early wet seasons. Pollen from nests collected in 1998–1999 without spatial information was also identified. 3. Bees used pollen of at least 64 species; many of these occurred in only one collection. The 2007 collections contained pollen of 35 different species, but were dominated by five species, especially Hura crepitans L. and Pseudobombax septenatum (Jacq.) Dugand. 4. Temporal availability, but not distance from nest, influenced flower use at a 50‐ha scale. 5. Body size was not associated with minimum flight distance as inferred from pollen collections. 6. Nest productivity and pollen diversity decreased from the dry to wet seasons, mirroring community‐level availability of floral resources. 7. Results suggest that on a scale of 50 ha, bees are choosing certain host plant species regardless of distance from the nest, but adjusting foraging behaviour opportunistically based on the temporal availability of host flowers.     

Intersexual mimicry and flowering phenology facilitate pollination in a dioecious habitat specialist species, <Emphasis Type="Italic">Myristica fatua</Emphasis> (Myristicaceae)

Myristica fatua is a dioecious specialist species restricted to the endangered, freshwater Myristica swamp forests in the Western Ghats, India. Earlier studies have alluded to pollination by deception in members of the Myristica genus, and thus we examined the pollination ecology comprising floral biology, flower production, flower visitors, and reproductive success in M. fatua and inferred the potential strategies that could permit such deception in this habitat specialist tree. Male flowers provide pollen rewards for an extended period of time while female flowers are rewardless and both sexes are visited by generalist insects, mainly by honeybees and stingless bees. Bee visits were significantly more frequent and longer on male than on female flowers as bees collected pollen from male flowers. We found that flower production patterns create a preponderance of males compared to females in the swamp populations. Using a model of honeybee color vision, we found the distance between the color loci of male and female flowers and based on minimum visual angle subtended by these flowers, we suggest that the two floral sexes cannot be discriminated by bees. Bees are likely deceived by the perceptual similarity of rewardless female flowers to pollen-offering male flowers and pollination is the consequence of foraging errors made by pollinators that encounter largely male–rarely female flower mosaics as they forage among clump-distributed M. fatua trees in the swamp habitat.     

The cuticular profiles of Australian stingless bees are shaped by resin of the eucalypt tree Corymbia torelliana

Bees are known to collect pollen and nectar to provide their larvae and themselves with food. That bees, especially the tropical stingless bees (Apidae: Meliponini), also collect plant resins has, however, been barely addressed in scientific studies on resource use in bees. Resins are used for nest construction, nest maintenance and nest defence. Furthermore, some South‐East Asian species transfer resin‐derived terpenes to their cuticular profiles. The resin requirement of bees is in turn used by certain plant species, which attract bees either for pollination by providing resin in their inflorescences, or for seed dispersal by providing resin in their seed capsules (mellitochory). Mellitochory is found in the eucalypt tree Corymbia torelliana, the resin of which is collected by Australian stingless bees. We investigated how the interaction between C. torelliana and resin‐collecting bees affects the chemical ecology of two Australian stingless bee genera by comparing the chemical profiles of eight bee species with resin from C. torelliana fruits. The two bee genera differed significantly in their chemical profiles. Similar to South‐East Asian stingless bees, 51% of all compounds on the body surfaces of the five Tetragonula species were most likely derived from plant resins. Up to 32 compounds were identical with compounds from C. torelliana resin, suggesting that Tetragonula species include C. torelliana compounds in their chemical profiles. By contrast, few or none resinous compounds were found on the body surfaces of the three Austroplebeia species sampled. However, one prominent but as yet unknown substance was found in both C. torelliana resin and the chemical profiles of all Tetragonula and four Austroplebeia colonies sampled, suggesting that most colonies (76%) gathered resin from C. torelliana. Hence, C. torelliana resin may be commonly collected by Australian stingless bees and, along with resins from other plant species, shape their chemical ecology.     

The influence of relative plant density and floral morphological complexity on the behaviour of bumblebees

We assessed the combined effects of varying the relative density and the relative floral morphological complexity of plant species on the behaviour of their bumblebee pollinators. Three species of bumblebee (Bombus pascuorum, B. terrestris and B. hortorum) were observed foraging on experimental arrays consisting of pair-wise combinations of four plant species: Borago officinalis, Phacelia tanacetifolia (both with simple flowers), Antirrhinum majus and Linaria vulgaris (both with complex flowers). Plant arrangements consisted of either two simple-flower species, a simple with a complex species or two complex species. The number of plants in each array was constant, while the frequency of each species was manipulated so that it was either rare, equal or common compared with its competitor. Contrary to predictions, rare plants were actually at an advantage in terms of the number of bees attracted per plant. However, rare plants were at a disadvantage in terms of pollen wastage because foragers more often went to a flower of another species after visiting a rare plant. The behaviour of bees on each plant species was further affected by plant floral complexity and the identity of the other species in the array. The three bumblebee species were markedly different in their foraging behaviour and in their responses to varying floral density and complexity. Each species preferred particular flower species. The results are discussed with reference to resource partitioning among bumblebee species. Received: 29 July 1998 / Accepted: 5 October 1998     

Development of a Species-Diagnostic Marker for Identification of the Stingless Bee Trigona pagdeni in Thailand

A species-diagnostic SCAR marker for identification of the stingless bee (Trigona pagdeni Schwarz) was successfully developed. Initially, amplified fragment length polymorphism analysis was carried out across representatives of 12 stingless bee species using 64 primer combinations. A 284 bp band restrictively found in T. pagdeni was cloned and sequenced. A primer pair (CUTP1-F/R) was designed and tested for species-specificity in 15 stingless bees. The expected 163 bp fragment was successfully amplified in all examined individuals of T. pagdeni (129/129). Nevertheless, cross-species amplification was also observed in T. fimbriata (1/3), T. collina (11/112), T. laeviceps (1/12), and T. fuscobalteata (15/15), but not in other species. SSCP analysis of CUTP1 further differentiated T. fuscobalteata and T. collina from T. pagdeni. Although T. laeviceps, T. fimbriata, and T. pagdeni shared an identical SSCP genotype, they are not taxonomically problematic species.     

Are pollination syndromes useful predictors of floral visitors in Tasmania?

Abstract Diurnal visitors to the flowers of many native plant species were identified in a wide range of Tasmanian sclerophyllous vegetation between September 1996 and April 1997. These foraging profiles were analysed to determine whether they were characteristic of various floral morphologies in predictable ways. It was found that although visitor profiles were sometimes consistent with classic pollination syndromes, these syndromes were unreliable predictors of floral visitors. Very few flowers were exclusively bird‐pollinated, and none were strictly fly‐, beetle‐, wasp‐, or butterfly‐pollinated. The majority of flowering plants were unspecialized in their morphology, and consequently hosted a diverse array of visitors. In addition, visitor profiles to congeners with similar floral morphologies, and even to conspecifics, differed between habitats. Altitude was a major factor in determining visitors, with flies being the most abundant visitors above 700 m. However, congeners in several genera of Epacridaceae, as well as the genus Correa, which differed in floral morphology also differed in visitor profiles. Tubular flowers were associated with birds, while flowers with more accessible nectar were visited by insects. The only taxa exhibiting a bee‐pollination syndrome that were largely visited by bees were the Fabaceae and Goodenia ovata Sm. Several species with purple or pink flowers were also predominantly visited by bees, but did not strictly conform to the melittophilous syndrome. In contrast, other flowers exhibiting an ostensibly mellitophilous syndrome hosted very few bees. Of these, species that occurred at high altitude were mainly visited by flies, while others received very few potential pollen vectors.     

Pollen partitioning of three species of Convolvulaceae among oligolectic bees in the Caatinga of Brazil

During the rainy season many species of Convolvulaceae bloom simultaneously in the Caatinga of northeast Brazil. In a Caatinga nature reserve we studied pollination and breeding systems of three sympatric species of Convolvulaceae, Ipomoea bahiensis, I.?nil, and Merremia aegyptia, focusing on pollen partitioning among flower visitors and pollen flow. The study showed that only oligolectic bees collected pollen and that these species had different preferences among the three species of Convolvulaceae: pollen of Ipomoea bahiensis, the only self-incompatible species, was collected mainly by Melitoma segmentaria, M.?osmioides, and Melitomella murihirta; pollen of I.?nil by Lithurgus huberi; and that of Merremia aegyptia by Ancyloscelis apiformis and an undescribed species of this genus. Introduced honey bees visited only flowers of Merremia aegyptia, where they were extraordinarily frequent flower visitors. However, they discarded the pollen grains, which led to almost 50% pollen loss. No polylectic bee species compete for pollen with the oligolectic species. Partitioning of pollen diminishes competition for floral resources in this specialized plant?Cpollinator association.     

Specialist <Emphasis Type="Italic">Osmia</Emphasis> bees forage indiscriminately among hybridizing <Emphasis Type="Italic">Balsamorhiza</Emphasis> floral hosts

Pollinators, even floral generalists (=polyleges), typically specialize during individual foraging bouts, infrequently switching between floral hosts. Such transient floral constancy restricts pollen flow, and thereby gene flow, to conspecific flowers in mixed plant communities. Where incipient flowering species meet, however, weak cross-fertility and often similar floral traits can yield mixed reproductive outcomes among pollinator-dependent species. In these cases, floral constancy by polyleges sometimes serves as an ethological mating barrier. More often, their foraging infidelities instead facilitate host introgression and hybridization. Many other bee species are oligolectic (taxonomic specialists for pollen). Oligoleges could be more discriminating connoisseurs than polyleges when foraging among their limited set of related floral hosts. If true, greater foraging constancy might ensue, contributing to positive assortative mating and disruptive selection, thereby facilitating speciation among their interfertile floral hosts. To test this Connoisseur Hypothesis, nesting females of two species of oligolectic Osmia bees were presented with randomized mixed arrays of flowers of two sympatric species of their pollen host, Balsamorhiza, a genus known for hybridization. In a closely spaced grid, the females of both species preferred the larger flowered B. macrophylla, evidence for discrimination. However, both species’ females showed no floral constancy whatsoever during their individual foraging bouts, switching randomly between species proportional to their floral preference. In a wider spaced array in which the bouquets reflected natural plant spacing, foraging oligolectic bees often transferred pollen surrogates (fluorescent powders) both between conspecific flowers (geitonogamy and xenogamy) and between the two Balsamorhiza species. The Connoisseur Hypothesis was therefore rejected. Foraging infidelity by these oligolectic Osmia bees will contribute to introgression and hybridization where interfertile species of Balsamorhiza meet and flower together. A literature review reveals that other plant genera whose species hybridize also attract numerous oligolectic bees, providing independent opportunities to test the generality of this conclusion.     

Recognition of scent marks in solitary bees to avoid previously visited flowers

Knowing how floral visitors forage efficiently among flowers is important to understanding plant-pollinator interactions. When bees search for rewarding flowers, they use several visual cues to detect the available floral resources. In addition to these cues, bees can recognize scent marks, which are olfactory cues left on flowers foraged by previous visitors. This behavior is well known in social bees, such as honeybees and bumblebees. Although solitary bees do not need to give information about which flowers were foraged to conspecifics, several pieces of evidence have indicated the use of scent marks. However, it is unknown whether the behavior is widely used in many different bee species. We investigated whether four different solitary bees, Colletes patellatus (Colletidae), Andrena prostomias (Andrenidae), Osmia orientalis (Megachilidae), and Tetralonia mitsukurii (Apidae), can recognize flowers that have been foraged previously by visitors within 3 min. All four bees showed rejection responses to flowers foraged by conspecifics. However, our results showed that responses to foraged flowers varied among bee species. The tendency of A. prostomias and T. mitsukurii to reject the foraged flowers was pronounced, while in C. patellatus and O. orientalis it was weak. In both A. prostomias and T. mitsukurii, the rejection rate of flowers foraged by conspecifics decreased as the time lag after the last visit increased. Both bees visited the flowers from which pollen or nectar had been artificially removed. We suggest that A. prostomias and T. mitsukurii would recognize scent marks left by previous visitors, while the other two bees would not recognize them so strongly. It is likely that the decision to use scent marks is dependent either on the richness of resources or on the complexity of floral structure.     

Pollen diets and niche overlap of honey bees and native bees in protected areas

The decline of both managed and wild bee populations has been extensively reported for over a decade now, with growing concerns amongst the scientific community. Also, evidence is growing that both managed and feral honey bees may exacerbate threats to wild bees. In Australia, there are over 1600 native bee species and introduced European honey bees (Apis mellifera) have established throughout most landscapes. There is a major gap in knowledge of the interactions between honey bees and native bees in Australian landscapes, especially floral resource use.Here we report on the pollen diets of wild bees in protected areas of coastal heathland, an ecosystem characterised by mass flowering in late winter and spring. We sampled bees within three sites and DNA metabarcoding was used to compare the pollen diets of honey bees and native bees. We recorded 2, 772 bees in total, with 13 genera and 18 described species identified. Apis mellifera was the most common species across all locations, accounting for 42% of all bees collected. Native bee genera included eusocial Tetragonula (stingless bees) (37%), and semi-social Exoneura and Braunsapis (19.8% combined). Metabarcoding data revealed both Tetragonula and honey bees have wide foraging patterns, and the bipartite network overall was highly generalised (H₂’ = 0.24). Individual honey bees carried pollen of 7–29 plant species, and significantly more species than all other bees. We found niche overlap in the diets of honey bees and native bees generally (0.42), and strongest overlap with stingless bees (0.70) and species of Braunsapis (0.62). A surprising finding was that many species carried pollen from Restionaceae and Cyperaceae, families generally considered to be predominantly wind-pollinated in Australia. Our study showed introduced honey bee use of resources overlaps with that of native bees in protected heathlands, but there are clear differences in their diet preferences.     

The pollination of the gynomonoecious Bdallophytum oxylepis (Cytinaceae,Malvales)

Bdallophytum oxylepis is a rare and endemic species belonging to the Cytinaceae family, a root holoparasitic plant in which most resources are allocated to attracting pollinators. This species is gynomonoecious with intraindividual variation in flower size and sex. Moreover, the flowers exhibit sapromyophilous traits, as do other species of Bdallophytum. Firstly, this study aimed to determine whether all floral morphs can form seeds and be pollen donors (in the case of bisexual flowers). Secondly, as this species has floral traits hypothesized to adapt to particular types of pollen vectors (carrion flies), we also studied the pollination of B. oxylepis to confirm whether the syndromes correspond to what occurs in nature. Through pollination treatments, we determined that all floral morphs are functional. By monitoring the inflorescences, we found that pollination is specialized in the studied population. Stingless bees performed pollination, as they have a high visitation rate, frequency, and constancy, and they are unique visitors that deposit pollen on the stigmas. Thus, they appear to be effective pollinators rather than carrion flies, as predicted by the syndrome. As shown here, animal–plant interaction studies can help establish a basis for conserving rare species such as holoparasites. Moreover, knowledge about the reproductive aspects of B. oxylepis reveals essential clues about its life cycle and role in maintaining native pollinators with economic and cultural value, such as stingless bees.     

Breeding system and pollination by mimicry of the orchid Tolumnia guibertiana in Western Cuba

The mimicry of malpighiaceous oil‐flowers appears to be a recurrent pollination strategy among many orchids of the subtribe Oncidiinae. These two plant groups are mainly pollinated by oil‐gathering bees, which also specialize in pollen collection by buzzing. In the present study, the floral ecology of the rewardless orchid Tolumnia guibertiana (Oncidiinae) was studied for the first time. The orchid was self‐incompatible and completely dependent on oil‐gathering female bees (Centris poecila) for fruit production. This bee species was also the pollinator of two other yellow‐flowered plants in the area: the pollen and oil producing Stigmaphyllon diversifolium (Malpighiaceae) and the polliniferous and buzzing‐pollinated Ouratea agrophylla (Ochnaceae). To evaluate whether this system is a case of mimetism, we observed pollinator visits to flowers of the three plant species and compared the floral morphometrics of these flowers. The behavior, preferences and movement patterns of Centris bees among these plants, as well as the morphological data, suggest that, as previously thought, flowers of T. guibertiana mimic the Malpighiaceae S. diversifolium. However, orchid pollination in one of the studied populations appears to depend also on the presence of O. agrophylla. Moreover, at the two studied populations, male and female pollination successes of T. guibertiana were not affected by its own floral display, and did not differ between populations. The results are discussed in relation to the behavior and preferences of Centris bees, as well as the differential presence and influence of each of the two floral models.     

Pollination of Campomanesia phaea (Myrtaceae) by night‐active bees: a new nocturnal pollination system mediated by floral scent

• Bees are the most important diurnal pollinators of angiosperms. In several groups of bees a nocturnal/crepuscular habit developed, yet little is known about their role in pollination and whether some plants are adapted specifically to these bees. We used a multidisciplinary approach to investigate the reproductive biology and to understand the role of nocturnal/crepuscular bees in pollination of Campomanesia phaea (Myrtaceae), popularly named cambuci.
• We studied the floral biology and breeding system of C. phaea. We collected the floral visitors and tested the pollinators' effectiveness. We also determined the floral scents released at night and during daytime, and studied behavioural responses of crepuscular/nocturnal bees towards these scents.
• The flowers of cambuci were self‐incompatible and had pollen as the only resource for flower visitors. Anthesis lasted around 14 h, beginning at 04:30 h at night. The flowers released 14 volatile compounds, mainly aliphatic and aromatic compounds. We collected 52 species of floral visitors, mainly bees. Nocturnal and crepuscular bees (four species) were among the most frequent species and the only effective pollinators. In field bioassays performed at night, nocturnal/crepuscular bees were attracted by a synthetic scent blend consisting of the six most abundant compounds.
• This study describes the first scent‐mediated pollination system between a plant and its nocturnal bee pollinators. Further, C. phaea has several floral traits that do not allow classification into other nocturnal pollination syndromes (e.g. pollinator attraction already before sunrise, with pollen as the only reward), instead it is a plant specifically adapted to nocturnal bees.

     

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.     

Initial floral visitor identity and foraging time strongly influence blueberry reproductive success

Priority effects occur when the order of species arrival affects subsequent ecological processes. The order that pollinator species visit flowers may affect pollination through a priority effect, whereby the first visitor reduces or modifies the contribution of subsequent visits. We observed floral visitation to blueberry flowers from honeybees, stingless bees or a mixture of both species and investigated how (i) initial visits differed in duration to later visits; and (ii) how visit sequences from different pollinator taxa influenced fruit weight. Stingless bees visited blueberry flowers for significantly longer than honeybees and maintained their floral visit duration, irrespective of the number of preceding visits. In contrast, honeybee visit duration declined significantly with an increasing number of preceding visits. Fruit weight was positively associated with longer floral visit duration by honeybees but not from stingless bee or mixed species visitation. Fruit from mixed species visits were heavier overall than single species visits, because of a strong priority effect. An initial visit by a stingless bee fully pollinated the flower, limiting the pollination contribution of future visitors. However, after an initial honeybee visit, flowers were not fully pollinated and additional visitation had an additive effect upon fruit weight. Blueberries from flowers visited first by stingless bees were 60% heavier than those visited first by honeybees when total floral visitation was short (∼1 min). However, when total visitation time was long (∼ 8 min), blueberry fruit were 24% heavier when initial visits were from honeybees. Our findings highlight that the initial floral visit can have a disproportionate effect on pollination outcomes. Considering priority effects alongside traditional measures of pollinator effectiveness will provide a greater mechanistic understanding of how pollinator communities influence plant reproductive success.