Co-dependency between a specialist <Emphasis Type="Italic">Andrena</Emphasis> bee and its death camas host, <Emphasis Type="Italic">Toxicoscordion paniculatum</Emphasis>

Among associations of plants and their pollinating bees, mutually specialized pairings are rare. Typically, either pollen specialist (oligolectic) bees are joined by polylectic bees in a flowering species’ pollinator guild, or specialized flowers are pollinated by one or more polylectic bees. The bee Andrena astragali is a narrow oligolege, collecting pollen solely from two nearly identical species of death camas (Toxicoscordion, formerly Zigadenus). Neurotoxic alkaloids of these plants are implicated in sheep and honey bee poisoning. In this study, T. paniculatum, T. venenosum and co-flowering forbs were sampled for bees at 15 sites along a 900-km-long east–west transect across the northern Great Basin plus an altitudinal gradient in northern Utah’s Bear River Range. Only A. astragali bees were regularly seen visiting flowering panicles of these Toxicoscordion. In turn, this bee was never among the 170 bee species caught at 17 species of other prevalent co-occurring wildflowers in the same five state region (38,000 plants surveyed). Our field pollination experiments show that T. paniculatum is primarily an outcrosser dependent on pollinator visitation for most capsule and seed set. Thus, both A. astragali and two sister species of Toxicoscordion are narrowly specialized and co-dependent on each other for reproduction, illustrating a rare case of obligate mutual specialization in bee–plant interactions.     

Thinking beyond Western commercial honeybee hives: towards improved conservation of honey bee diversity

A decline of wild pollinators, along with a decline of bee diversity, has been a cause of concern among academics and governmental organizations. According to IPBES, a lack of wild pollinator data contributes to difficulties in comprehensively analyzing the regional status of wild pollinators in Africa, Latin America, Asia and Oceania. It may have also contributed to the prevailing lack of awareness of the diversity of honey bees, of which the managed Apis mellifera is often considered as “the (only) honey bee,” despite the fact that there are eight other honey bee species extant in Asia. A survey of 100 journal articles published in 2016 shows that 57% of the studies still identified A. mellifera as “the honey bee.” In total, 80% of studies were conducted solely on A. mellifera. This focus on A. mellifera has also caused the honey standard of Codex Alimentarius and the European Union to be based solely on A. mellifera, causing improper evaluation of honeys from other species. We recommend adapting current standards to reflect the diversity of honey bees and in the process correct failures in the honey market and pave the way towards improved protection of honey bee species and their habitats.     

Non-native plants affect generalist pollinator diet overlap and foraging behavior indirectly,via impacts on native plant abundance

Flowering invasive plants can have dramatic effects on the resource landscape available to pollinators. Because many pollinators exhibit behavioral plasticity in response to competitor or resource density, this in turn can result in impacts on ecological processes such as pollination and plant reproduction. We examine how interactions between five common generalist eusocial bees change across an invasion gradient by examining how bee abundance and diet overlap changed with variation in both invasive plant abundance and competitor abundance in a temperate oak-savannah ecosystem. Specifically we focus on the bumblebees Bombus bifarius, B. mixtus, B. melanopygus and B. vosnesenskii, as well as the non-native honeybee Apis mellifera, and their interactions with the native flowering plants Camassia quamash, Camassia liechtlinii, and the invasive shrub Cytisus scoparius. We further examine whether changes in pollinator visits to the invasive and two common native plants can explain changes in diet overlap. Abundance of the invasive plant and other common floral resources had strong impacts on focal bee abundance, with certain species more likely to be present at highly invaded sites. This may be because highly invaded sites tended to be embedded in forested landscapes where those bees are common. Diet overlap was most affected by abundance of a common native plant, rather than the invasive plant, with diet overlap increasing non-linearly with abundance of the native plant. Furthermore, Apis mellifera, did not appear to have direct competitive effects on native bumblebees in this habitat. However, visit patterns suggest that bees most abundant at highly invaded sites may compete for access to native resources. Thus the impacts of this invasive plant on our focal bee species may be primarily indirect, via its’ competitive effects on native plants.     

Age-related changes in the behavioural response of honeybees to <Emphasis Type="Italic">Apiguard</Emphasis><Superscript>®</Superscript>, a thymol-based treatment used to control the mite <Emphasis Type="Italic">Varroa destructor</Emphasis>

The parasitic mite Varroa destructor is responsible for heavy losses in honey bee colonies and represents a major threat to the beekeeping industry. Essential oils offer an attractive alternative to the use of synthetic chemicals for the control of varroa. Amongst them, thymol appears to be particularly promising. However, treatments using thymol as their active substance, such as the gel formulation Apiguard ^®, are suspected to have adverse effects on honey bee colonies. In this study, laboratory assays are used to investigate the effects of Apiguard ^® exposure on honey bee behaviour. Our results reveal that honey bee responses to this anti-varroa treatment change with honey bee age. While 2-day-old bees respond neutrally to Apiguard ^®, older bees generally avoid the Apiguard ^® gel. Responses of forager bees were particularly striking. Foragers appear to be repelled by Apiguard ^®. Touching their antennae with Apiguard ^® induces robust fanning behaviour. Our data suggest, however, that forager bees exposed to Apiguard ^® in the hive can become habituated to this treatment. These results offer interesting new perspectives on the effects of Apiguard ^® on honey bee behaviour and serve to highlight age-related changes in honey bee responses to gustatory, as well as olfactory cues.     

Bee visitors of <Emphasis Type="Italic">Centaurea solstitialis</Emphasis> L. (Asteraceae) in an urban environment in northwestern Turkey

Information on the pollination ecology and floral visitors of the noxious weed Centaurea solstitialis is available for several populations in its invasive range, but limited information is available in its native range, with most studies conducted on the Greek island of Lesvos. Herein, we document the visitation pattern of bees and explore the relationship of bee body size and nectar availability in weedy populations of C. solstitialis from an urban environment within its native range in northwestern Turkey. Studies were conducted at patches of C. solstitialis in abandoned lots at the Uluda? University near the city of Bursa. A total of 41 species, including honey bees, belonging to five families and 19 genera were recorded. Small megachilid and halictid bee species were the most common visitors. Average nectar standing crop volume per floret was low (0.003–0.117 μL) and did not significantly vary throughout the day. Average bee head width significantly correlated with average nectar standing crop volume but did not significantly change throughout the day. Analyses of pollen loads as well as direct observations of bee foraging behavior indicate that a large number of bees visit C. solstitialis, primarily in search of nectar while carrying a large percentage of pollen grains of this plant species on their bodies. These results are similar to previous observations on a non-weedy population of C. solstitialis from the island of Lesvos.     

Novel microsatellite loci reveal high genetic diversity yet low population structure for alfalfa leafcutting bees in North America

The alfalfa leafcutting bee, Megachile rotundata (ALCB) is an economically important pollinator necessary for seed production of the critical forage crop alfalfa, Medicago sativa. The pollinator was accidentally introduced to North America from Europe approximately 70 years ago, and it is primarily produced in Canada and shipped to the United States annually en masse for seed field pollination. We investigate how the large-scale commercial movement of this bee affects the genetic structure of populations in the North American seed growing system and compare the genetic diversity and structure of introduced North American bees with two native European populations. Using 16 newly developed microsatellite loci, we describe the North American population structure of this bee. ALCBs collected from alfalfa seed farms have a degree of genetic variability similar to one native European population, but lower than the second. Considering that the species was accidentally introduced into North America, we anticipated more signature of a founder effect. Despite the level of genetic variability, we found little, if any, genetic structuring across North America, other than that the North American populations were distinct from the European populations sampled. While we detected some sub-structure in North American populations using Bayesian methods, the structuring was without geographic pattern, and we propose it is the result of the intense human management and movement of these bees. The trade and movement of these bees by humans has created a nearly panmictic M. rotundata population across the continent, which has implications relevant to the preservation and conservation of other bee pollinators.     

More than protein? Bee–flower interactions and effects of disturbance regimes revealed by rare pollen in bee nests

Bees and their host flower populations were studied by identifying pollen to species or genus, from trap nests where bees were reared. Rare plant species in bee diets, and disturbance regimes, have not previously been researched and are emphasized here. Two focal bee groups with one species each (Megachilidae and Apidae) were studied in a 500,000-ha tropical reserve in the Yucatán Peninsula nine complete years. The number of rare or major pollen species in nests had no statistical correlation; thus, rare pollen analysis complements study of major brood provisions. We found most nests (87 % Megachile zaptlana, 93 % Centris analis) contained rare pollen; only 12 % of the 438 nests contained major pollen alone. Rare pollen sometimes indicated an energy source rather than a scarce protein resource. Trichome nectar of Cydista, along with Ipomoea and Caesalpinia, were nectar sources. Malpighiaceae, despite lacking nectar, often provided the complete Centris diet. Considering rare pollen, only Centris responded to drought, or competition from immigrant honeybees. Neither bee responded to hurricanes. Drought years coincided with low bee populations; Centris nests contained more rare species then. After feral Africanized honeybees colonized, Centris had more major species and fewer rare. Some herbarium vouchers from the study area contained exotic pollen, demonstrating in situ floral contamination and ecological generalization by bees, but this rarely occurred in plants found among the bee diets. Megachile and Centris responded differently to competition and resource scarcity, and plausibly evolved under different disturbance regimes, yet appeared well adapted to hurricane disturbance.     

The origins of global invasions of the German wasp (<Emphasis Type="Italic">Vespula germanica</Emphasis>) and its infection with four honey bee viruses

A successful control or eradication programme using biological control or genetically-mediated methods requires knowledge of the origin and the extent of wasp genetic diversity. Mitochondrial DNA variation in the native and invaded range of the social wasp Vespula germanica was used to examine intra-specific genetic variation and invasive source populations. We also examined wasps for the presence of four viruses found in honey bees: Acute bee paralysis virus, Deformed wing virus, Israeli acute paralysis virus and Kashmir bee virus. German wasps showed reduced genetic diversity in the invaded range compared to that of their native range. Populations in the introduced range are likely to have arrived from different source populations. All four viral honey bee pathogens were found in V. germanica, although they varied in their distribution and strain. Multiple introductions of German wasps have occurred for most invaded regions, though some populations are genetically homogenous. The differing locations of origin will guide researchers searching for biocontrol agents and the reduced genetic diversity may make these wasps a potentially viable target for control via gene drives.     

Influence of landscape context on the abundance of native bee pollinators in tomato crops in Central Brazil

Evidence shows that pollinator abundance has declined and, consequently, so has their services, which has possible negative impacts on ecosystem functioning. The goal of this study was to evaluate the influence of landscape context at multiple spatial scales on the abundance of bee pollinators of tomato crops in Brazil. Pollinator abundance was obtained from tomato crops grown in a conventional system in the Cerrado region. Around each tomato field circular buffers of 0.75, 1, 1.5, 2, and 3 km radius were defined. Inside each buffer the landscapes were manually classified into native and non-native cover and, the proportion of native vegetation, the relative largest patch size, and the distance of the nearest native vegetation to each field were calculated. Pollinator species were categorized into five groups: Exomalopsis, Centris, Bombus/Eulaema, Halictidae, and all buzz pollinators combined (Buzzers). The results showed that the landscape context influenced the abundance of the five groups of tomato pollinators. Bees with a smaller body size, such as Exomalopsis spp., responded at smaller scales, while bees with a larger body size, such as the Centris and Bombus/Eulaema groups, responded at larger scales. The abundance of all pollinator groups increased with native vegetation cover. Most groups showed higher abundances in landscapes with similar-size fragments. The results reinforce the recommendation for maintaining natural habitats around crop areas, even if fragmented, for the conservation of the tomato pollinator assemblage. These findings are valuable for planning landscape management in the studied area to improve bee conservation, ecosystem services, and food production.     

Pollinator-mediated impacts of alien invasive plants on the pollination of native plants: the role of spatial scale and distinct behaviour among pollinator guilds

Alien invasive plant species can affect pollination, reproductive success and population dynamics of co-flowering native species via shared pollinators. Consequences may range from reproductive competition to facilitation, but the ecological drivers determining the type and magnitude of such indirect interactions remain poorly understood. Here, we examine the role of the spatial scale of invader presence and spatially contingent behavioural responses of different pollinator groups as potential key drivers, using the invasive Oxalis pes-caprae and the self-incompatible native annual Diplotaxis erucoides as a model system. Three treatments were assigned to native focal plants: (1) invader present at the landscape scale (hectares) but experimentally removed at the floral neighbourhood scale (pa); (2) invader present at both scales (pp); (3) invader absent at both scales (aa). Interestingly, we found pronounced spatially contingent differences in the responses of pollinators: honeybees and bumblebees were strongly attracted into invaded sites at the landscape scale, translating into native plant visitation facilitation through honeybees, while bumblebees almost exclusively visited Oxalis. Non-corbiculate wild bees, in contrast, showed less pronounced responses in foraging behavior, primarily at the floral neighborhood scale. Average heterospecific (Oxalis) pollen deposition onto stigmas of Diplotaxis was low (<1 %), but higher in the pp than in the pa treatment. Hand-pollination of Diplotaxis with Oxalis and conspecific pollen, however, reduced seed set by more than half when compared to hand-pollination with only conspecific pollen. Seed set of Diplotaxis, finally, was increased by 14 % (reproductive facilitation) in the pp treatment, while it was reduced by 27 % (reproductive competition) in the pa treatment compared to uninvaded populations. Our study highlights the crucial role of spatial scale and pollinator guild driving indirect effects of invasive on co-flowering native plant species.     

Floral divergence and temporal pollinator partitioning in two synchronopatric species of <Emphasis Type="Italic">Vigna</Emphasis> (Leguminosae-Papilionoideae)

Exclusivity of pollinators, temporal partitioning of shared pollinators and divergence in pollen placement on the shared pollinators’ bodies are mechanisms that prevent interspecific pollen flow and minimize competitive interactions in synchronopatric plant species. We investigated the floral biology, flower visitors, pollinator effectiveness and seasonal flower availability of two syntopic legume species of the genus Vigna, V. longifolia and V. luteola, in ‘restinga’ vegetation of an island in southern Brazil. Our goal was to identify the strategies that might mitigate negative consequences of their synchronous flowering. Vigna longifolia and V. luteola were self-compatible, but depended on pollinators to set seeds. Only medium to large bees were able to trigger the ‘brush type’ pollination mechanism. Vigna longifolia, with its asymmetrical corolla and hugging mechanism, showed a more restrictive pollination system, with precise sites of pollen deposition/removal on the bee’s body, compared to V. luteola, with its zygomorphic corolla and cymbiform keel. There was a daily temporal substitution in flower visitation by the main pollinators. Vigna longifolia and V. luteola had overlapping flowering phenology but the densities of their flowers fluctuated, resulting in a seasonal partitioning of flower visitation. The differences in corolla symmetry and mainly the temporal partitioning among pollinators throughout the day and the flowering season proved to be important factors in maintaining the synchronopatry of V. longifolia and V. luteola.     

Hunting for healthy microbiomes: determining the core microbiomes of <Emphasis Type="Italic">Ceratina,Megalopta</Emphasis>, and <Emphasis Type="Italic">Apis</Emphasis> bees and how they associate with microbes in bee collected pollen

Social corbiculate bees such as honey bees and bumble bees maintain a specific beneficial core microbiome which is absent in wild bees. It has been suggested that maintaining this microbiome can prevent disease and keep bees healthy. The main aim of our study was to identify if there are any core bacterial groups in the non-corbiculate bees Ceratina and Megalopta that have been previously overlooked. We additionally test for associations between the core bee microbes and pollen provisions to look for potential transmission between the two. We identify three enterotypes in Ceratina samples, with thirteen core bacterial phylotypes in Ceratina females: Rosenbergiella, Pseudomonas, Gilliamella, Lactobacillus, Caulobacter, Snodgrassella, Acinetobacter, Corynebacterium, Sphingomonas, Commensalibacter, Methylobacterium, Massilia, and Stenotrophomonas, plus 19 in pollen (6 of which are shared by bees). Unlike Apis bees, whose gut microbial community differs compared to their pollen, Ceratina adults and pollen largely share a similar microbial composition and enterotype difference was largely explained by pollen age. Megalopta displays a highly diverse composition of microbes throughout all adults, yet Lactobacillus and Saccharibacter were prevalent in 90% of adults as core bacteria. Only Lactobacillus was both a core bee and pollen provision microbe in all three species. The consequences of such diversity in core microbiota between bee genera and their associations with pollen are discussed in relation to identifying potentially beneficial microbial taxa in wild bees to aid the conservation of wild, understudied, non-model bee species.     

Introgression in native populations of <Emphasis Type="Italic">Apis mellifera mellifera</Emphasis> L: implications for conservation

Hybridisation and introgression can have negative impacts on regional biodiversity through the potential erosion of locally adapted lineages. The honey bee (Apis mellifera L.) occurs in twenty-seven subspecies across Europe, is an extremely economically important insect, yet threatened by multifarious impacts. Transhumance of the most commercially appealing varieties threatens native honey bee diversity by introgression and subsequent loss of locally adapted traits, or even by complete removal of some subspecies from parts of the range. Here levels of admixture and introgression are examined in UK honey bees suspected to be from hives of the dark European honey bee (Apis mellifera mellifera). Microsatellite DNA and STRUCTURE analyses reveal that the studied populations are generally admixed, and discriminant analysis of principal components shows them to be intermediate between A. m. mellifera and Apis mellifera carnica populations. However, examining mitochondrial haplotype data (COI-COII intergenic spacer region) and nuclear DNA reveal that some hives are relatively pure (from 4 to 15 hives, depending on the Q-value threshold). Genetic diversity is relatively high in comparison with other European populations. Implications for conservation and management are discussed.     

Population genetics of wild and managed pollinators: implications for crop pollination and the genetic integrity of wild bees

Loss of habitat and chemical use associated with agriculture can cause population declines of wild pollinators. Less is known about the evolutionary consequences of interactions between species used in commercial agriculture and wild pollinators. Given population declines of many wild bee species, it is crucial to understand if commercial queens become established in natural areas, if wild bees visit agricultural fields and have the potential to interact with commercial bees, and if gene flow occurs between commercial and wild bees. We drew on a long-term data set that documents commercial bumble bee (Bombus impatiens) use in New England, and we conducted genetic analyses of foraging B. impatiens from areas with varying intensities of commercial bee use. In agricultural areas with a history of commercial bee use we also sampled bees directly from commercial hives. We found significant genetic differences among foraging B. impatiens and B. impatiens sampled directly from hives (average pairwise F′_ST = 0.14), but not among samples of foraging bees from natural areas (average F′_ST among foraging bees?=?0.002). Furthermore, Bayesian analysis of population structure revealed that foraging bees caught in areas with a history of commercial bee use grouped with samples from natural areas. These results document an agricultural setting where there was no widespread introgression of alleles from commercial bumble bees to wild bumble bees, commercial bumble bees did not become established in natural areas, and wild bees were providing pollination services to crops.     

Pollinators and crossability as reproductive isolation barriers in two sympatric oil-rewarding <Emphasis Type="Italic">Calceolaria</Emphasis> (Calceolariaceae) species

Pollinator species are widely accepted as an important factor in plant reproductive isolation. Although mostly investigated in plants visited by different groups of pollinators (e.g., hummingbirds vs bees), few studies have examined the role of pollinators belonging to the same taxonomic group (e.g., only bees) on plant reproductive isolation. In this study, we investigate this question by evaluating pre- and post-zygotic mechanisms putatively involved in the reproductive isolation of two oil-rewarding sympatric Calceolaria species (i.e., Calceolaria filicaulis and C. arachnoidea) in an Andean ecosystem of Chile. We estimated reproductive isolation values using a combination of field (pollinator visitation rates) and experimental (intra and interspecific manual cross-pollination and seed germination of parents and hybrids) evidence. The two Calceolaria species were preferentially visited by different oil-collecting bee species. Results from hand cross-pollination experiments indicate that intraspecific crossings produced significantly more seeds than interspecific ones. Notwithstanding, seed germination essays did not reveal differences between parental and hybrids. Taken together, these results suggest that pollinator species are responsible for most of the reproductive isolation in the two Calceolaria species studied here. This study is the first assessment of pollinator-mediated reproductive isolation in Calceolaria species and the first to document reproductive barriers in oil-rewarding plants.     

Native bees provide insurance against ongoing honey bee losses

One of the values of biodiversity is that it may provide 'biological insurance' for services currently rendered by domesticated species or technology. We used crop pollination as a model system, and investigated whether the loss of a domesticated pollinator (the honey bee) could be compensated for by native, wild bee species. We measured pollination provided to watermelon crops at 23 farms in New Jersey and Pennsylvania, USA, and used a simulation model to separate the pollen provided by honey bees and native bees. Simulation results predict that native bees alone provide sufficient pollination at > 90% of the farms studied. Furthermore, empirical total pollen deposition at flowers was strongly, significantly correlated with native bee visitation but not with honey bee visitation. The honey bee is currently undergoing extensive die-offs because of Colony Collapse Disorder. We predict that in our region native bees will buffer potential declines in agricultural production because of honey bee losses.     

Pollen load diversity and foraging niche overlap in a pollinator community of the rare <Emphasis Type="Italic">Dictamnus albus</Emphasis> L.

Bees collect pollen as an important resource for offspring development while acting as pollen vectors for the plants visited. Foraging preferences of pollinators together with plant species availability shape the web of interactions at the local scale. In this study, we focused on the bee pollinator community of a population of the rare protected perennial herb Dictamnus albus, with the aim to characterise the pollen preferences and the foraging niche overlap among species through time. Bees were sampled during four consecutive years in a natural population of D. albus, throughout the blooming period of the plant species. We performed an analysis of insect pollen loads to investigate the interactions with the study species and the co-flowering plants in the area, and to evaluate the degree of foraging overlap among pollinators. Over the study years, all bee species showed a high fidelity to D. albus (60–80%), even if some taxa preferentially collected pollen from other flowering species. The foraging niche overlap in the pollinator community decreased together with an increased diversity of co-flowering plant species. The results obtained indicate that bees preferentially forage on D. albus in the studied area, but that co-flowering species contribute to complement their diet and likely reduce competition for foraging resources. It appears therefore important to maintain a high diversity of co-flowering plants to preserve the diversity in the studied pollinator community of D. albus.     

Multitrophic interactions among fungal endophytes,bees, and <Emphasis Type="Italic">Baccharis dracunculifolia</Emphasis>: resin tapering for propolis production leads to endophyte infection

The tropics are known for their high diversity of plants, animals, and biotic interactions, but the role of the speciose endophytic fungi in these interactions has been mostly neglected. We report a unique interaction among plant sex, bees, and endophytes on the dioecious shrub, Baccharis dracunculifolia (Asteraceae). We assessed whether there was an association between resin collection by bees and fungal endophytes considering the host plant sex. We hypothesized that resin collection by the Africanized honey bee, Apis mellifera L. (Apidae) could favor the entry of endophytes in B. dracunculifolia. Specifically, we tested the hypotheses that (1) bees damage the leaf buds of female and male plant at different proportions; (2) damage on leaf buds increases the richness of endophytic fungi; (3) endophyte richness differs between female and male plants; and (4) in vitro growth of endophytes depends on the sex of the plant individual from which the resin was extracted. Endophyte richness and proportion of leaf bud damage did not vary between the plant sexes. However, species similarity of endophytes between female and male plants was 0.33. Undamaged leaf buds did not show culturable endophytes, with all fungi exclusively found in damaged leaf buds. Endophyte composition changed with the plant sex. The endophytes exclusively found in female plants did not develop in the presence of male resin extract. These findings highlight that resin collection by A. mellifera for propolis production favors the entry of endophytic fungi in B. dracunculifolia. Additionally, endophyte composition and growth are influenced by plant sex.     

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.     

Increase in ant density promotes dual effects on bee behaviour and plant reproductive performance

Floral rewards do not only attract pollinators, but also herbivores and their predators. Ants are attracted by extrafloral nectaries (EFNs), situated near flowers, and may interfere with the efficiency and behaviour of pollinators. We tested the hypothesis that the impacts of ant–pollinator interactions in plant–pollinator systems are dependent on (1) the seasonal activity of EFNs, which increase ant abundance closer to flowers; (2) consequently, an ant effect, where ants decrease the temporal niche overlap of bees due to predator avoidance; and (3) ant density, where higher densities may negatively affect plant–pollinator interactions and plant performance. We studied two ant–plant–pollinator systems based on Banisteriopsis campestris and Banisteriopsis malifolia plant species. The periods of high ant abundance coincided with plant species blooming. The presence of ants around flowers reduced the visitation rates of the smaller bees and the temporal niche overlap between bee species was not higher than randomly expected when ants had free access. Additionally, we observed variable ant effects on fruit set and duration of bee visits to both Malpighiaceae species when ant density was experimentally kept constant on branches, especially on B. campestris. Our goal was to show the dual role of ant density effects, especially because the different outcomes are not commonly observed in the same plant species. We believe that reduced temporal niche overlap between floral visitors due to ant presence provides an opportunity for smaller bees to improve compatible pollination behaviour. Additionally, we concluded that ant density had variable effects on floral visitor behaviours and plant reproductive performance.