In flowers with poricidal anthers, pollen is not freely accessible and legitimate access is restricted to bees capable of vibrating the anthers. Despite the protection of pollen provided by poricidal anthers, numerous illegitimate, non‐buzz‐pollinating flower visitors rob pollen. We aimed to quantify the influence of functional groups of floral visitors and illegitimate interactions on the network structure to disentangle the flower visitor network into its mutualistic and antagonistic components. We delimited three functional groups of bees based on their pollen collection behaviour in poricidal flowers: large bees that vibrate entire flowers in a single buzzing position (flower buzzing), bees vibrating single anthers in different positions (anther buzzing) and non‐vibrating flower‐damaging or gleaning bees (non‐buzzing). Moreover, we characterized legitimate and illegitimate interactions of co‐occurring and co‐flowering plants and their flower visitors based on the stigma contact during a visit. Since we independently assessed the type of interaction with bee–plant species combinations, we were able to include the behavioural variations of each bee species across different flowers. The networks were modular, with stronger interactions within subsets of species than among the subsets. All modules included a combination of flower‐, anther‐ and non‐buzzing bees, and mutualistic and antagonistic networks were intermingled. Seven bee species shifted their roles across plant species. Specialization in the subset of interactions with pollinators was higher than the overall visitation network. Flower‐buzzing bees were more specialized than anther‐buzzing and non‐buzzing bees, which used virtually all poricidal flowers similarly. Although plants with poricidal anthers shared a specialized mechanism of pollen release, their pollinators were highly dissimilar and formed compartments of interacting species. The interaction‐level approach taken in our study confers a high specificity to the pollinator network, leading to a more complex and realistic picture of mutualistic webs versus its embedded florivory, which are otherwise confounded in pooled networks across flower visitors. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13204/suppinfo is available for this article.

Heteranthery, the presence of feeding and pollinating anthers in the same flower, seems to mediate the evolutionary dilemma for plants to protect their gametes and yet provide food for pollinators. This study aims to elucidate the role of heteranthery in the buzz-pollinated Senna reniformis. The fecundity of pollen from long-, medium- and short-sized anthers was determined by hand cross-pollination experiments, and the quantity, size, ornamentation and viability of pollen of different anthers were compared. Rates of flower rejection by bees were measured in anther removal experiments to assess the preferences of flower visitors for feeding or pollinating anthers. Large bees, which were the effective pollinators of self-incompatible S. reniformis, avoided flowers without short feeding anthers, but not those without medium or long anthers. Illegitimate small and medium-sized bees were unresponsive to anther exclusion experiments. Long anthers deposited pollen on the back and short anthers on the venter of large bees. Pollen from long anthers had higher in vitro viability and higher fruit and seed set after cross-pollination than pollen from other sized anthers. Short anthers produce feeding pollen to effective pollinators and long anthers are related to pollination of S. reniformis. Bee behaviour and size was found to directly influence the role of anthers in the 'division of labour'. Only large bee pollinators that carry the pollinating pollen from long anthers in 'safe sites' associated short anthers with the presence of food. In the absence of these larger bee pollinators, the role of heteranthery in S. reniformis would be strongly compromised and its function would be lost.

PREMISE Flowering plants with poricidal anthers are commonly visited by buzzing bees, which vibrate flowers to extract pollen. However, not all flower visitors are in fact pollinators, and features such as body size and duration of flower visits are important factors in determining pollination effectiveness. We tested whether bee‐to‐flower size relationships predict the pollination effectiveness of flower visitors of a buzz‐pollinated species (Chamaecrista ramosa, Fabaceae). METHODS We sorted 13 bee taxa into three groups: smaller than, equivalent to (“fit‐size”), and larger than flower herkogamy (spatial separation between anthers and stigma). We expected the latter two groups to touch the stigmas, which would be an indicator of pollination effectiveness, more frequently than the first group. To test this hypothesis, we assessed contact with stigmas, foraging behavior, and duration of visits for the three size groups of bees. RESULTS Our data reveal that small bees scarcely touched the stigmas, while large and fit‐size bees were the most efficient pollinators, achieving high stigma‐touching rates, conducting much shorter flower visits, and visiting flowers and conspecific plants at high rates during foraging bouts. CONCLUSIONS The results did not show size‐matching among bees and flowers, as expected, but rather a minimum size threshold of efficient pollinators. The finding of such a threshold is a nonarbitrary approach to predicting pollination effectiveness of visitors to herkogamous flowers with poricidal anthers.

ResumoA leguminosa Leucaena leucocephala (Lam.) R. de Wit., conhecida como leucena, é originária da América Central, considerada pioneira, com populações de altas densidades, o que dificulta o desenvolvimento das espécies nativas. O presente estudo objetivou contribuir com informações sobre a biologia reprodutiva da L. leucocephala, referentes ao seu êxito reprodutivo. O estudo foi realizado em Trindade, Goiás, de julho a novembro de 2012. Foram selecionadas quatro áreas de estudo no município apresentando populações da leucena constituídas por, no mínimo, seis indivíduos em plena floração. Foram realizadas observações dos visitantes florais e testes de primeira, segunda e terceira visitas, além de polinização cruzada entre plantas, autopolinização e controle. Foram registrados 342 visitantes, distribuídos em 12 espécies de insetos com predominância de Apis mellifera (Linnaeus, 1758) e Trigona spinipes (Fabricius, 1793) nas áreas. A flor apresentou senescência ao fim do dia, mas houve decréscimo na abundância de insetos nos indivíduos de leucena após as 10 horas da manhã. Na avaliação do número de visitas de polinizadores e sua influência na produção de frutos, verificou-se que o número de visitas do principal polinizador (A. mellifera) não influenciou a produção de frutos, evidenciando a eficiência da abelha desde a primeira visita na flor. Leucaena leucocephala reúne vários atributos típicos de uma espécie invasora, como a ação eficiente de polinizadores generalistas (A. mellifera), propiciando produção considerável de frutos e sementes, contribuindo para o potencial dispersor da planta e sucesso reprodutivo nas áreas.Palavras-chave: polinização, Apis mellifera, sucesso reprodutivo. AbstractThe species Leucaena leucocephala (Lam.) R. de Wit., known as leucaena, is originally from Central America. It has been considered a pioneer with high densities of individuals in its populations that could prevent the development of native species. The present study aims to understand the reproductive biology of L. leucocephala, synthetizing information about its reproductive success. The study was conducted in Trindade, central Brazil, from July to November 2012. All areas contained specimens in bloom. We selected four study areas containing populations of leucaena with at least six individuals with flowering. Observations were made of floral visitors and we did tests of first, second and third visit, and cross-pollination between plants, self-pollination and control group (without human intervention). There was a predominance of Apis mellifera and Trigona spinips as flower visitors in these areas. The flowers undergo senescence at the end of the day, however, there is a decrease in flower visiting by insects after 10 am. We found 342 visitors belonging to 12 insect species. After assessing the number of visits of pollinators and their influence on fruit production, we found that the number of visits of the main pollinator (A. mellifera) does not influence the production of fruits, showing the efficiency of this bee in its...

Background Chile is the second largest avocado producer in the world. However, there have been few studies on interactions between native bees and avocado pollination in a Chilean environment. The flowering period of avocado is very sensitive to temperature fluctuations and so environmental conditions, particularly cool temperatures, can have a strong influence on its floral behaviour. Thus, we aimed to determine whether temperature also influences bee visitation rates to avocado flowers or if this was due to the number of available flowers. Moreover, we proposed to verify the attractiveness of avocado flowers to native bees and compare flower handling time and type of floral resource collected between managed and native bees. Methods An avocado orchard of “Hass” cultivar located in the Mediterranean region of Chile was studied during the flowering season of 2016. The frequency of exotic and native bee species in flowers, floral phenology, and environmental temperature were analysed, and the foraging behaviours of exotic and native bee species in avocado flowers compared. Results Five Chilean native and two exotic species were recorded visiting avocado flowers at the studied orchard. The bees generally sought nectar rather than pollen. The European honeybee (Apis mellifera) was the most frequent flower visitor, followed by the native species Colletes cyanescens and Cadeguala occidentalis. There was a strong positive correlation between temperature and number of visits, but no significant correlation between number of open flowers and number of visits. Conclusions Native Chilean bees visited avocado flowers and some of them were frequent floral visitors. They generally sought for nectar rather than pollen and were as fast as honeybees handling avocado flowers. The environmental temperature plays a key role in the frequency of bees in avocado flowers. The warmer weather is related to the higher number of bees visits to avocado flowers.

1. Blueberry is one of the most relevant buzz-pollinated crops worldwide and Chile is the most important global producer of fresh blueberries during wintertime in the Northern Hemisphere. Thousands of exotic Bombus terrestris are imported from Europe to pollinate blueberries. However, no study has investigated the performance of the native Chilean fauna to pollinate blueberry or other crops. Therefore, we aimed to compare the performance of native Chilean floral visitors with managed visitors to pollinate highbush blueberry.2. Per-visit pollination performance (stigmatic pollen deposition) and floral visitation were measured and the presence of sonication behaviour of flower visitors was evaluated for five cultivars in two blueberry orchards located in southern Chile.3. Floral visitors showed a preference for one or more blueberry cultivars, instead of visiting all cultivars equally. Floral visits with sonication deposited more conspecific pollen on stigmas than visits without sonication. Some native sonicating bees (Cadeguala and Bombus), especially Cadeguala occidentalis, were efficient pollen vectors of blueberry and better pollinators than honeybees (5.8 times more pollen transferred) similar to that of the managed bee B. terrestris. 4.The results indicate that some Chilean native bee species, especially those with sonication behaviour, can provide pollination service to highbush blueberry crops. ResumenAbejas nativas con comportamiento de sonicaci on floral pueden lograr un alto rendimiento de la polinizaci on de arándano alto en Chile

The specialised mutualism between Tococa guianensis and ants housed in its leaf domatia is a well-known example of myrmecophily. A pollination study on this species revealed that flowers in the bud stage exude a sugary solution that is collected by ants. Given the presence of this unexpected nectar secretion, we investigated how, where, and when floral buds of T. guianensis secret nectar and what function it serves. We studied a population of T. guianensis occurring in a swampy area in the Cerrado of Brazil by analyzing the chemical composition and secretion dynamics of the floral-bud nectar and the distribution and ultrastructure of secretory tissues. We also measured flower damage using ant-exclusion experiments. Floral bud nectar was secreted at the tip of the petals, which lack a typical glandular structure but possess distinctive mesophyll due to the presence of numerous calcium oxalate crystals. The nectar, the production of which ceased after flower opening, was composed mainly of sucrose and low amounts of glucose and fructose. Nectar was consumed by generalist ants and sporadically by stingless bees. Ant exclusion experiments resulted in significantly increased flower damage. The floral nectar of T. guianensis is produced during the bud stage. This bud-nectar has the extranuptial function of attracting generalist ants that reduce florivory. Pollen is the unique floral resource attracting pollinators during anthesis. Tococa guianensis, thus, establishes relationships with two functional groups of ant species: specialist ants acting against herbivory and generalist ants acting against florivory.

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