Pollen dispensing schedules in buzz-pollinated plants: experimental comparison of species with contrasting floral morphologies.

PREMISE: Plants can mitigate the fitness costs associated with pollen consumption by floral visitors by optimizing pollen release rates. In buzz-pollinated plants, bees apply vibrations to remove pollen from anthers with small pores. These poricidal anthers potentially function as mechanism staggering pollen release, but this has rarely been tested across plant species differing in anther morphology. METHODS: In Solanum Section Androceras, three pairs of buzz-pollinated species have undergone independent evolutionary shifts between large- and small-flowers, which are accompanied by replicate changes in anther morphology. We used these shifts in anther morphology to characterize the association between anther morphology and pollen dispensing schedules. We applied simulated bee-like vibrations to anthers to elicit pollen release, and compared pollen dispensing schedules across anther morphologies. We also investigated how vibration velocity affects pollen release. RESULTS: Replicate transitions in Solanum anther morphology are associated with consistent changes in pollen dispensing schedules. We found that small-flowered taxa release their pollen at higher rates than their large-flowered counterparts. Higher vibration velocities resulted in quicker pollen dispensing and more total pollen released. Finally, both the pollen dispensing rate and the amount of pollen released in the first vibration were negatively related to anther wall area, but we did not observe any association between pore size and pollen dispensing. CONCLUSIONS: Our results provide the first empirical demonstration that the pollen dispensing properties of poricidal anthers depend on both floral characteristics and bee vibration properties. Morphological modification of anthers could thus provide a mechanism to exploit different pollination environments.

High species richness of native pollinators in Brazilian tomato crops / Alta riqueza de espécies de polinizadores nativos em plantações Brasileiras de tomateiro

Abstract Pollinators provide an essential service to natural ecosystems and agriculture. In tomatoes flowers, anthers are poricidal, pollen may drop from their pore when flowers are shaken by the wind. However, bees that vibrate these anthers increase pollen load on the stigma and in fruit production. The present study aimed to identify the pollinator richness of tomato flowers and investigate their morphological and functional traits related to the plant-pollinator interaction in plantations of Central Brazil. The time of anthesis, flower duration, and the number and viability of pollen grains and ovules were recorded. Floral visitors were observed and collected. Flower buds opened around 6h30 and closed around 18h00. They reopened on the following day at the same time in the morning, lasting on average 48 hours. The highest pollen availability occurred during the first hours of anthesis. Afterwards, the number of pollen grains declined, especially between 10h00 to 12h00, which is consistent with the pollinator visitation pattern. Forty bee species were found in the tomato fields, 30 of which were considered pollinators. We found that during the flowering period, plants offered an enormous amount of pollen to their visitors. These may explain the high richness and amount of bees that visit the tomato flowers in the study areas. The period of pollen availability and depletion throughout the day overlapped with the bees foraging period, suggesting that bees are highly effective in removing pollen grains from anthers. Many of these grains probably land on the stigma of the same flower, leading to self-pollination and subsequent fruit development. Native bees (Exomalopsis spp.) are effective pollinators of tomato flowers and are likely to contribute to increasing crop productivity. On the other hand, here tomato flowers offer large amounts of pollen resource to a high richness and amount of bees, showing a strong plant-pollinator interaction in the study agroecosystem.

Resumo Polinizadores fornecer um serviço essencial para os ecossistemas naturais e para agricultura. Em tomateiros, as anteras são poricidas e o pólen pode sair a partir dos poros quando as flores são agitadas pelo vento. No entanto, as abelhas que vibram as anteras aumentam a carga de pólen no estigma e na produção de frutos. O presente estudo teve como objetivo identificar a riqueza dos polinizadores das flores de tomate e investigar suas características morfológicas e funcionais relacionadas com a interação planta-polinizador em plantações do Brasil central. Foram registrados o tempo de antese, duração flor, bem como o número e viabilidade de grãos de pólen e óvulos. Os visitantes florais foram observados e coletados. Os botões florais abriram-se em torno 06h30 e fechou em torno de 18h00. As flores reabrem no dia seguinte ao mesmo tempo na parte da manhã, com longevidade média de 48 horas. A maior disponibilidade de pólen ocorreu durante as primeiras horas da antese. Depois disso, o número de grãos de pólen diminuiu, especialmente entre as 10h00 às 12h00, o que é consistente com os padrões de visitação de polinizadores. Quarenta espécies de abelhas foram encontradas nos campos de tomate, 30 das quais foram consideradas polinizadores. Durante o período de floração, as plantas oferecem enorme quantidade de pólen para os seus visitantes. Isto pode explicar a alta riqueza e quantidade de abelhas que visitam as flores de tomate nas áreas de estudo. O período de disponibilidade e redução de pólen durante todo o período do dia é sobreposto com o período de alimentação das abelhas, o que sugere que elas são altamente eficazes na remoção de grãos de pólen das anteras. Muitos desses grãos provavelmente são depositados no estigma da mesma flor, levando à auto-polinização e o desenvolvimento de frutos. Abelhas nativas (Exomalopsis spp.) são polinizadores efetivos de flores de tomate, podendo contribuir para o aumento da produtividade das culturas. Por outro lado, as flores de tomate oferecem grandes quantidades de pólen de recursos para uma alta riqueza e quantidade de abelhas, que mostram um estudo forte interação planta-polinizador nos agroecossistemas.

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

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

Abelhas (Apoidea) visitantes de flores de Comolia ovalifolia DC Triana (Melastomataceae) em uma área de restinga na Bahia / Bee (Apoidea) visitors of the flowers of Comolia ovalifolia DC Triana (Melastomataceae) in a sandbanks area in Bahia

The flowers of Comolia ovalifolia present poricidal anthers and their pollen serve as protein source to a specific group of bees, able to extract the pollen from the anthers through vibration. From January to December /2001, we observed the flowering of C. ovalifolia and the activity of the most frequent bees throughout the year. The peaks of flowering of C. ovalifolia occurred in March and from September to November/2001. The flowers of C. ovalifolia were visited by 16 species of bees, being the most frequent Centris leprieuri (Spinola), Xylocopa subcyanea Pérez, Centris sp., Euglossa sp. and Florilegus similis Urban. Among these, C. leprieuri was the most frequent visitor in the flowers (89%) and its foraging period overlapped with the blooming of C. ovalifolia. This bee presented adequate size and efficient behavior in pollen collection and transfer to the stigma, being considered an effective pollinator of the flowers.

As flores de Comolia ovalifolia apresentam anteras poricidas e seu pólen serve como fonte protéica a um grupo específico de abelhas, capaz de extrair o pólen das anteras através de vibração. O florescimento de C. ovalifolia e a atividade das abelhas mais freqüentes foram observadas de janeiro a dezembro/2001. Os picos de florescimento de C. ovalifolia ocorreram nos meses de março/2001 e entresetembro e novembro/2001. As flores de C. ovalifolia foram visitadas por 16 espécies de abelhas, sendo as mais freqüentes Centris leprieuri (Spinola), Xylocopa subcyanea Pérez, Centris sp., Euglossa sp. e Florilegus similis Urban. Entre elas, C. leprieuri destacou-se com 89% das visitas e sua atividade forrageira anual coincidiu com a floração de C. ovalifolia. Além disso, essa abelha apresentou tamanho adequado e comportamento eficiente na coleta e transferência de pólen para o estigma, sendo considerada polinizador legítimo dessas flores.