The invasive bumblebee Bombus terrestris (Linnaeus, 1758) disrupts the adaptive function of heteranthery by indiscriminately visiting the pollinating and feeding anthers of Senna arnottiana flowers.

Heteranthery, the presence of different types of anthers on the same flower, is a floral adaptation that aims to balance the need for pollinators to collect pollen as a food resource while ensuring sufficient pollen for pollination. We investigate the role of heteranthery in the pollination of Senna arnottiana flowers and how it is affected by the behaviour of visiting bee species, with a particular focus on the impact of the invasive bumblebee Bombus terrestris. In three populations of S. arnottiana we measured the size of three sets of anthers and style, stigma-anther separation, pollen quantity and fruit set, and contrasted it with the body size, behaviour, and pollination effectiveness of all floral visitors. Different bee species visited S. arnottiana flowers, and their foraging behaviour varied. Large-bodied native bees, including Centris cineraria, Caupolicana sp. and Cadeguala occidentalis, preferentially visited short anthers, whereas B. terrestris, an exotic bumblebee, foraged from both short and long anthers without distinction. In addition, B. terrestris contacted the stigma at a lower rate than large-bodied native bees. Instead of concentrating its pollen-gathering efforts on the feeding anthers, as predicted by the "division of labor" hypothesis, B. terrestris indiscriminately visited both types of anthers similarly. This behaviour of B. terrestris may disrupt the adaptive significance of heteranthery by mixing the roles of pollination and feeding anthers of S. arnottiana. Therefore, our results highlight the potential disruption of this relationship by exotic pollinators and the need to consider it in conservation efforts.

Biomechanical properties of non-flight vibrations produced by bees.

Bees use thoracic vibrations produced by their indirect flight muscles for powering wingbeats in flight, but also during mating, pollination, defence and nest building. Previous work on non-flight vibrations has mostly focused on acoustic (airborne vibrations) and spectral properties (frequency domain). However, mechanical properties such as the vibration's acceleration amplitude are important in some behaviours, e.g. during buzz pollination, where higher amplitude vibrations remove more pollen from flowers. Bee vibrations have been studied in only a handful of species and we know very little about how they vary among species. In this study, we conducted the largest survey to date of the biomechanical properties of non-flight bee buzzes. We focused on defence buzzes as they can be induced experimentally and provide a common currency to compare among taxa. We analysed 15,000 buzzes produced by 306 individuals in 65 species and six families from Mexico, Scotland and Australia. We found a strong association between body size and the acceleration amplitude of bee buzzes. Comparison of genera that buzz-pollinate and those that do not suggests that buzz-pollinating bees produce vibrations with higher acceleration amplitude. We found no relationship between bee size and the fundamental frequency of defence buzzes. Although our results suggest that body size is a major determinant of the amplitude of non-flight vibrations, we also observed considerable variation in vibration properties among bees of equivalent size and even within individuals. Both morphology and behaviour thus affect the biomechanical properties of non-flight buzzes.

Heteromorphic stamens are differentially attractive in Swartzia (Fabaceae).

The division of labour hypothesis between stamens has explained the evolution of divergent functions between dimorphic stamens in the same flower. However, little is known about whether the distinct type of stamens differs in attractiveness to pollinators. Therefore, we investigate whether the two types of stamens commonly found in Swartzia have different visual and olfactory attractants. We performed observations of anthesis dynamics, registration and collection of floral visitors, measurements of reflectance of floral parts and chemical analysis of the volatile organic compounds of the floral parts of two species, S. flaemingii and S. simplex. Both species have two distinct sets of stamens: one with smaller and abundant stamens in the centre of the flower and the other with fewer but larger abaxial stamens. The sets differ in UV reflectance (only S. simplex) and exhibit a distinct chromatic contrast. Concerning olfactory attractiveness, aliphatic compounds make up most of the odour of the two species, both whole flowers and most of their floral organs. On the other hand, only S. simplex presented apocarotenoids (as ionones) and benzenoids. Furthermore, there are differences in the proportion of volatiles emitted by the stamen in both cases, as the high proportion of sesquiterpenes among the smaller stamens compared to the larger ones. In conclusion, the two types of stamens found in S. flaemingii and S. simplex show a distinct attractiveness. In addition, our data have demonstrated diverse ways of differential attractiveness both between distinct stamens set per flower and between the two species from the same pollen flowers genus.

A sophisticated case of division of labour in the trimorphic stamens of the Cassia fistula (Leguminosae) flower.

Buzz-pollinated pollen flowers have pollen as the primary resource for pollinators and must deal with a conflict between the exploitation of pollen grains by bees and pollination success. It has been hypothesized that heterostemony allows division of labour between stamens as a solution to the pollen dilemma. To test the division of labour hypothesis, we chose Cassia fistula, which has a trimorphic androecium and analysed androecium development, pollen grain release mechanisms and visitor behaviour. We explored the reflectance of floral organs and carried out an exclusion experiment to test the attractiveness of each stamen morph to the bee species. Finally, we explored the structural, ultrastructural and functional variation between the pollen grains, including pollen viability across stamen morphs. The differences among the three stamen morphs, which is developed from two whorls of the stamen, are the first evidence of the division of labour in our study system. Large Bombus and Xylocopa bees actively and exclusively exploited the pollen grains from the central poricidal anthers generating pollen deposition on their bodies. The reflectance pattern of floral organs indicated a targeting of these large bees to the central anthers, corroborated by the anther manipulative experiment where only the exclusion of the anthers positioned in the flower centre, especially the intermediate stamens, reduced bee visits. Both results revealed a division of labour, in which the intermediate stamen morph was responsible for both floral attractiveness and pollen resources. Only the largest stamen morph produced germinable pollen grains, highlighting their role as pollinating stamens. The smallest stamen morph has a less clear function, likely representing an economy in pollen production for feeding function. Our findings suggest that the evolution of the trimorphic androecium is associated with division of labour in large pollen flowers and can represent a strong strategy for circumventing the pollen dilemma, optimizing the feeding function by reducing pollen grain investment from central anthers.

Minimum size threshold of visiting bees of a buzz-pollinated plant species: consequences for pollination efficiency.

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.

Connective appendages in Huberia bradeana (Melastomataceae) affect pollen release during buzz pollination.

Floral structures, such as stamen appendages, play crucial roles in pollinator attraction, pollen release dynamics and, ultimately, the reproductive success of plants. The pollen-rewarding, bee buzz-pollinated flowers of Melastomataceae often bear conspicuous staminal appendages. Surprisingly, their functional role in the pollination process remains largely unclear. We use Huberia bradeana Bochorny & R. Goldenb. (Melastomataceae) with conspicuously elongated, twisted stamen appendages to investigate their functional role in the pollination process. We studied the effect of stamen appendages on pollinator behaviour and reproductive success by comparing manipulated flowers (appendages removed) with unmanipulated flowers. To assess bee pollinator behaviour, we measured three properties of buzzes (vibrations) produced by bees on Huberia flowers: frequency, duration and number of buzzes per flower visit. We measured male and female reproductive success by monitoring pollen release and deposition after single bee visits. Finally, we used artificial vibrations and laser vibrometry to assess how flower vibrational properties change with the removal of stamen appendages. Our results show that the absence of staminal appendages does not modify bee buzzing behaviour. Pollen release was higher in unmanipulated flowers, but stigmatic pollen loads differ only marginally between the two treatments. We also detected lower vibration amplitudes in intact flowers as compared to manipulated flowers in artificial vibration experiments. The presence of connective appendages are crucial in transmitting vibrations and assuring optimal pollen release. Therefore, we propose that the high diversity of colours, shapes and sizes of connective appendages in buzz-pollinated flowers may have evolved by selection through male fitness.

Bee-Mediated Selection Favors Floral Sex Specialization in a Heterantherous Species: Strategies to Solve the Pollen Dilemma.

Animal-pollinated plants show a broad variation in floral morphology traits and gametophyte production within populations. Thus, floral traits related to plant reproduction and sexuality are usually exposed to pollinator-mediated selection. Such selective pressures may be even stronger in heterantherous and pollen flowers, in which pollen contributes to both bee feeding and pollination, overcoming the "pollen dilemma" or the inability to perform both functions simultaneously. We describe the phenotypic gender and sexual organ morphology of flowers in two populations of Macairea radula (Melastomataceae), a heterantherous and buzz-pollinated species with pollen flowers. We estimated selection gradients on these traits through female and male fitness components. Both populations showed sizeable phenotypic gender variation, from strict hermaphrodites to increased femaleness or maleness. We found a continuous variation in style and stamen size, and this variation was correlated with corresponding shape values of both sexual organs. We detected bee-mediated selection towards short and long styles through seed number and towards intermediate degrees of heteranthery through pollen removal in one population, and selection towards increased maleness through pollen dispersal in both populations. Our results suggest that bee-mediated selection favors floral sex specialization and stylar dimorphism in M. radula, optimizing reproductive success and solving the pollen dilemma.

Heteranthery as a solution to the demand for pollen as food and for pollination - Legitimate flower visitors reject flowers without feeding anthers.

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.

Ricochet pollination in Senna (Fabaceae) - petals deflect pollen jets and promote division of labour among flower structures.

Naturalists Fritz and Hermann Müller hypothesised that heteranthery often leads to a division of labour into 'feeding' and 'pollinating' stamens; the latter often being as long as the pistil so as to promote successful pollination on the bees' back. In many buzz-pollinated species of Senna, however, the so-called pollinating stamens are short and not level with the stigma, raising the question of how pollen is shed on the bees' back. Here we explore a mechanism called 'ricochet pollination'. We test whether division of labour is achieved through the interaction between short lower stamens and strongly concave 'deflector petals'. We studied the arrangement and morphology of the floral organs involved in the ricochet pollination, functioning of the flowers through artificial sonication and observed the interactions between bees and flowers in the field. The middle stamens are adapted to eject pollen downwards, which can be readily collected on the bee mid legs. Most of the pollen is ejected towards the deflector petal(s). Pollen from this set of stamens is more likely to contribute to pollination. The pollen grains seem to ricochet multiple times against the deflector petals to eventually reach the bee's back. The pollen ricochet mechanism promotes a division of labour by involving additional floral organs, such as petals, reinforcing the Müllers' division-of-labour hypothesis. However, alternative, non-multiexclusive hypotheses could be explored in genus Senna and other angiosperm species.

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.

High species richness of native pollinators in Brazilian tomato crops

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.

Apomixis does not affect visitation to flowers of Melastomataceae, but pollen sterility does.

Apomixis is an asexual seed reproduction mechanism thorough which embryos are originated from material tissues inside the ovules, without precedent fertilisation. It allows plants to colonise new habitats, even in places where flower visitors are scarce or where plants are isolate. Apomixis seems to be related to pollen sterility and, in species with flowers that offer pollen as a reward for pollinators, the amount or quality of the pollen offered by these species may influence the amount of the visits and specific composition of the visitors. In order to test this hypothesis, we studied breeding systems of 16 species of Melastomataceae and their flower visitors, evaluating composition and abundance of the visits to apomictic and sexual species. Apomictic plants with no viable pollen or with pollen with low viability did not receive visits from pollinators, and consequently probably produce strictly apomictic fruits. On the other hand, apomictic and sexual plants with high pollen viability do receive visits; in this case, apomictic plants may produce fruits and seeds through both sexual and apomictic methods. The species composition of insects visiting Melastomataceae with high pollen viability was similar, regardless of whether the plants were apomictic or not. It seems that pollen viability levels are important to determine visits to the flowers irrespective of breeding system.

Biología floral y reproductiva de Vaccinium meridionale (Ericaceae) en los Andes orientales de Colombia

Vaccinium meridionale es una planta silvestre productora de frutos comestibles en las áreas montañosas del norte de Suramérica. Sin embargo, no hay información suficiente sobre su sistema reproductivo, lo cual es fundamental para su manejo y conservación dado que sus frutos son extraídos de forma no sostenible y hay un creciente interés en establecer la especie como cultivo. Los objetivos de este trabajo fueron estudiar la biología floral y reproductiva de V. meridionale en condiciones naturales y analizar la importancia de los polinizadores en su reproducción. Para esto se realizaron observaciones y experimentos para describir su morfología y fenología floral, viabilidad de polen, receptividad de estigma, relación polen óvulo y producción de néctar. Para estudiar su sistema reproductivo se realizaron experimentos de emasculación, exclusión de polinizadores y polinización manual (autopolinización y polinización cruzada). El estudio se realizó con dos poblaciones silvestres de V. meridionale ubicadas en los departamentos de Cundinamarca y Boyacá, en la Cordillera Oriental de Colombia. Se encontró que aunque las flores tienen anteras poricidas la liberación del polen se da fácilmente sin vibración. V. meridionale presenta un gran despliegue y larga longevidad floral y hay un sesgo en la producción de néctar hacia la función femenina. Se estimó una relación polen/óvulo de 571±133, lo que clasifica a la especie como xenógama facultativa. Este resultado coincidió con los experimentos de polinización ya que en las dos poblaciones estudiadas las plantas produjeron frutos por agamospermia, autogamia y xenogamia. No obstante, se observó una fuerte depresión por endogamia manifestada en altas tasas de aborto de frutos producidos por autogamia a diferencia de los frutos provenientes de polinización cruzada que fueron retenidos por la planta desde su formación. Los rasgos florales presentados por la especie se consideran mecanismos para favorecer un gremio más diverso de visitantes florales que sólo los insectos capaces de polinización por vibración. Además, estos rasgos florales pueden aumentar la probabilidad de polinización y disminuir la geitonogamia. Igualmente la depresión por endogamia sugiere que la especie busca mantener el entrecruzamiento como su principal estrategia reproductiva. Por lo tanto, los polinizadores, en particular las abejas son fundamentales para la reproducción de V. meridionale y en consecuencia su conservación es esencial para mantener la variabilidad genética y oferta de frutos de esta especie.

Vaccinium meridionale is a wild plant producing edible fruits in the mountain areas of Northern South America. However, the fruits of this species has been under an unsustainable extraction and there is a growing interest of establishing this species as a crop; nevertheless, the information about its breeding system is scarce, which is essential for its sustainable management and conservation. This research aimed to study the floral and reproductive biology of V. meridionale in natural conditions, and to analyze the importance of pollinators on its reproduction, in two wild populations of V. meridionale in the states of Cundinamarca and Boyacá, in the Oriental Cordillera of Colombia. For this, we have made different observations and experiments to describe its flower morphology, floral phenology, pollen viability, stigma receptivity, pollen-ovule ratio and nectar production. To study its reproductive system, we performed experiments of flower emasculation, pollinator exclusion and hand pollination (self-and cross-pollination). We found that although the flowers have poricidal anthers, the release of pollen could occur easily without vibration. V. meridionale shows a large floral display, long floral longevity and has female-biased nectar production. The pollen-ovule ratio was of 571±133, which classified the species as facultative xenogamy. This result agreed with the pollination experiments because the plants produced fruits by agamospermy, selfing and outcrossing. However, we registered a strong inbreeding depression, observed in high rates of fruit abortions, after self-pollination. Unlike of self-pollinating fruits, the plant retains those produced by cross-pollination since its formation. The floral traits showed by this species are mechanisms to favor a more diverse guild of floral visitors than only insects able to buzz-pollination. In addition, these floral traits may enhance the pollination probability, and reduce geitonogamy. Moreover, the inbreeding depression suggests that V. meridionale promotes outcrossing as its main reproductive strategy. Therefore, pollinators, particularly bees, are essential for this species reproduction and conservation, and are critical in the maintenance of its genetic variability and fruits production.

Potential pollinators of Comolia ovalifolia DC Triana (Melastomataceae) and Chamaecrista ramosa (Vog.) H.S. Irwin and Barneby var. ramosa (Leguminosae-Caesalpinioideae), in restinga, Bahia, Brazil / Polinizadores potenciais de Comolia ovalifolia DC Triana (Melastomataceae) e Chamaecrista ramosa (Vog.) H.S. Irwin e Barneby var. ramosa (Leguminosae-Caesalpinioideae), na restinga, Bahia, Brasil

Comolia ovalifolia DC Triana (Melastomataceae) and Chamaecrista ramosa (Vog.) H.S. Irwin and Barneby var. ramosa (Leguminosae - Caesalpinioideae) are tropical plant species found in restinga (herbaceous-shrubby, sandy costal ecosystems). They have flowers with poricidal anthers and are pollinated by bees. The study sought to analyse potential pollinators of both plants during visits to their flowers in a restinga area in Bahia. The flowering displayed by both species was considered continuous and long duration, constantly providing pollen to floral visitors. C. ovalifolia was visited by 17 species of bees and C. ramosa by 16 species, predominantly from the Apidae family (with a similarity index of 74 percent). The behavior displayed by these visiting bees was of vibrating anthers. The small-sized Euglossa sp. Latreille, 1802 and Florilegus similis Urban, 1970 bees played less of a role as pollinators, since they rarely touched the flower stigma during harvests and were thus considered opportunist visitors or casual pollinators. Centris decolorata Lepetier, 1841 (= C. leprieuri) and Xylocopa subcyanea Perez, 1901 are large bees and were considered efficient pollinators of C. ovalifolia and C. ramosa because of the higher frequency and constancy of their visits, and their favourable behaviour and size for pollen transfer between flowers, which guarantees the survival of these native restinga plant species.

Comolia ovalifolia (Melastomataceae) e Chamaecrista ramosa (Leguminosae - Caesalpinioideae) são espécies de plantas tropicais que ocorrem na restinga. Estas plantas apresentam flores com anteras poricidas e são polinizadas por abelhas. Este estudo teve como objetivo analisar os polinizadores potenciais de ambas as plantas durante as visitas nas flores em uma área de restinga da Bahia. O florescimento apresentado por ambas as espécies foi considerado contínuo e de longa duração, oferecendo sempre pólen para os visitantes florais. C. ovalifolia foi visitada por 17 espécies de abelhas e C. ramosa por 16 espécies, com predominância da família Apidae (índice de similaridade de 74 por cento). As abelhas visitantes destas flores apresentavam o comportamento de vibrar as anteras. O papel das abelhas de pequeno porte como Euglossa sp. e Florilegus similis, como polinizadores, era pequeno, pois raramente tocavam o estigma das flores durante as coletas, sendo consideradas visitantes oportunistas ou polinizadores casuais. Centris decolorata e Xylocopa subcyanea, são abelhas grandes, e foram consideradas polinizadores eficientes de C. ovalifolia e C. ramosa, por apresentarem elevada frequência e constância de visitas nestas plantas, e comportamento e porte favoráveis à transferência do pólen entre as flores, o que garante a sobrevivência destas espécies de plantas nativas da restinga.

Potential pollinators of Comolia ovalifolia DC Triana (Melastomataceae) and Chamaecrista ramosa (Vog.) H.S. Irwin and Barneby var. ramosa (Leguminosae-Caesalpinioideae), in restinga, Bahia, Brazil / Polinizadores potenciais de Comolia ovalifolia DC Triana (Melastomataceae) e Chamaecrista ramosa (Vog.) H.S. Irwin e Barneby var. ramosa (Leguminosae-Caesalpinioideae), na restinga, Bahia, Brasil

Comolia ovalifolia DC Triana (Melastomataceae) and Chamaecrista ramosa (Vog.) H.S. Irwin and Barneby var. ramosa (Leguminosae - Caesalpinioideae) are tropical plant species found in restinga (herbaceous-shrubby, sandy costal ecosystems). They have flowers with poricidal anthers and are pollinated by bees. The study sought to analyse potential pollinators of both plants during visits to their flowers in a restinga area in Bahia. The flowering displayed by both species was considered continuous and long duration, constantly providing pollen to floral visitors. C. ovalifolia was visited by 17 species of bees and C. ramosa by 16 species, predominantly from the Apidae family (with a similarity index of 74 percent). The behavior displayed by these visiting bees was of vibrating anthers. The small-sized Euglossa sp. Latreille, 1802 and Florilegus similis Urban, 1970 bees played less of a role as pollinators, since they rarely touched the flower stigma during harvests and were thus considered opportunist visitors or casual pollinators. Centris decolorata Lepetier, 1841 (= C. leprieuri) and Xylocopa subcyanea Perez, 1901 are large bees and were considered efficient pollinators of C. ovalifolia and C. ramosa because of the higher frequency and constancy of their visits, and their favourable behaviour and size for pollen transfer between flowers, which guarantees the survival of these native restinga plant species.(AU)

Comolia ovalifolia (Melastomataceae) e Chamaecrista ramosa (Leguminosae - Caesalpinioideae) são espécies de plantas tropicais que ocorrem na restinga. Estas plantas apresentam flores com anteras poricidas e são polinizadas por abelhas. Este estudo teve como objetivo analisar os polinizadores potenciais de ambas as plantas durante as visitas nas flores em uma área de restinga da Bahia. O florescimento apresentado por ambas as espécies foi considerado contínuo e de longa duração, oferecendo sempre pólen para os visitantes florais. C. ovalifolia foi visitada por 17 espécies de abelhas e C. ramosa por 16 espécies, com predominância da família Apidae (índice de similaridade de 74 por cento). As abelhas visitantes destas flores apresentavam o comportamento de vibrar as anteras. O papel das abelhas de pequeno porte como Euglossa sp. e Florilegus similis, como polinizadores, era pequeno, pois raramente tocavam o estigma das flores durante as coletas, sendo consideradas visitantes oportunistas ou polinizadores casuais. Centris decolorata e Xylocopa subcyanea, são abelhas grandes, e foram consideradas polinizadores eficientes de C. ovalifolia e C. ramosa, por apresentarem elevada frequência e constância de visitas nestas plantas, e comportamento e porte favoráveis à transferência do pólen entre as flores, o que garante a sobrevivência destas espécies de plantas nativas da restinga.(AU)

Reproductive system of Eriocnema fulva Naudin (Melastomataceae), an endemic species of Minas Gerais state, SE Brazil

Eriocnema fulva Naudin is a perennial herb, endemic to Minas Gerais state, SE Brazil, found on humid, shaded rocky riverbanks in montane semideciduous seasonal forests. The species is threatened, but information regarding its biology is still lacking, although such information is fundamental to any management plan. We aimed to evaluate the reproductive system of Eriocnema fulva in the Jambreiro Forest (19° 58'-59' S and 43° 55'-52' W, 800-1100 m altitude), municipality of Nova Lima, by experiments carried out in 1997 and 1998. The flowers are white, and flowering is of the steady state type, occurring once a year from November to December. Anthers are poricidal, and pollen is the only resource for visitors. The chromosome number is n = 17 during meiosis. The species is self-compatible, but does not produce fruits by spontaneous self-pollination or agamospermy; it requires pollen vectors and buzz pollination in order to produce fruits. Cross-pollination is the main reproductive strategy of E. fulva, and is accentuated by the small number of flowers (one or two in each plant) opened per day. Although the population studied was shaded by forest canopy, the seeds needed light to germinate. Germination ratio was lower in germination cabinet on filter paper (14 percent after 30 days) than in greenhouse on soil brought from the forest (47 percent after 25 days). Although the fruit is a capsule and the seeds are small, dispersion (anemochory or hydrochory) does not seem to occur at long distance, as it is the case for other Melastomataceae species with similar syndrome.

Eriocnema fulva Naudin é uma planta herbácea, perene, endêmica no estado de Minas Gerais, sendo encontrada em paredões rochosos úmidos, ao longo de riachos sombreados pela Floresta Estacional Semidecídua Montana. A espécie é ameaçada de extinção e não existem informações sobre a sua biologia, embora sejam fundamentais para um plano de manejo. Com o objetivo de avaliar o sistema de reprodução, foram feitos experimentos em 1997 e 1998 em uma população na Mata do Jambreiro (19° 58'-59' S e 43° 55'-52' W, 800-1100 m de altitude), no município de Nova Lima. As flores são brancas e a floração é do tipo steady state, ocorrendo uma vez ao ano durante novembro e dezembro. As anteras são poricidas e o pólen é a única recompensa para os visitantes. O número cromossômico encontrado na meiose foi n = 17. A espécie é autocompatível, não produz frutos por autopolinização espontânea nem por agamospermia, mas requer obrigatoriamente vetores de pólen e polinização vibrátil. A polinização cruzada é a principal estratégia reprodutiva de Eriocnema fulva, sendo acentuada devido ao pequeno número de flores abertas por dia, apenas uma ou duas em cada planta. Embora as populações estejam localizadas em ambientes sombreados e úmidos, as sementes precisam de luz para germinar, sendo fotoblásticas positivas. A taxa de germinação em papel de filtro na câmara de germinação foi menor (14 por cento após 30 dias) que em solo da floresta na casa de vegetação (47 por cento após 25 dias). Embora o fruto seja do tipo cápsula com sementes pequenas, a dispersão (anemocoria ou hidrocoria) parece não ocorrer a distâncias longas, como é observada em outras espécies de Melastomataceae com síndrome de dispersão semelhante.

Reproductive system of Eriocnema fulva Naudin (Melastomataceae), an endemic species of Minas Gerais state, SE Brazil

Eriocnema fulva Naudin is a perennial herb, endemic to Minas Gerais state, SE Brazil, found on humid, shaded rocky riverbanks in montane semideciduous seasonal forests. The species is threatened, but information regarding its biology is still lacking, although such information is fundamental to any management plan. We aimed to evaluate the reproductive system of Eriocnema fulva in the Jambreiro Forest (19° 58'-59' S and 43° 55'-52' W, 800-1100 m altitude), municipality of Nova Lima, by experiments carried out in 1997 and 1998. The flowers are white, and flowering is of the steady state type, occurring once a year from November to December. Anthers are poricidal, and pollen is the only resource for visitors. The chromosome number is n = 17 during meiosis. The species is self-compatible, but does not produce fruits by spontaneous self-pollination or agamospermy; it requires pollen vectors and buzz pollination in order to produce fruits. Cross-pollination is the main reproductive strategy of E. fulva, and is accentuated by the small number of flowers (one or two in each plant) opened per day. Although the population studied was shaded by forest canopy, the seeds needed light to germinate. Germination ratio was lower in germination cabinet on filter paper (14% after 30 days) than in greenhouse on soil brought from the forest (47% after 25 days). Although the fruit is a capsule and the seeds are small, dispersion (anemochory or hydrochory) does not seem to occur at long distance, as it is the case for other Melastomataceae species with similar syndrome.

Eriocnema fulva Naudin é uma planta herbácea, perene, endêmica no estado de Minas Gerais, sendo encontrada em paredões rochosos úmidos, ao longo de riachos sombreados pela Floresta Estacional Semidecídua Montana. A espécie é ameaçada de extinção e não existem informações sobre a sua biologia, embora sejam fundamentais para um plano de manejo. Com o objetivo de avaliar o sistema de reprodução, foram feitos experimentos em 1997 e 1998 em uma população na Mata do Jambreiro (19° 58'-59' S e 43° 55'-52' W, 800-1100 m de altitude), no município de Nova Lima. As flores são brancas e a floração é do tipo steady state, ocorrendo uma vez ao ano durante novembro e dezembro. As anteras são poricidas e o pólen é a única recompensa para os visitantes. O número cromossômico encontrado na meiose foi n = 17. A espécie é autocompatível, não produz frutos por autopolinização espontânea nem por agamospermia, mas requer obrigatoriamente vetores de pólen e polinização vibrátil. A polinização cruzada é a principal estratégia reprodutiva de Eriocnema fulva, sendo acentuada devido ao pequeno número de flores abertas por dia, apenas uma ou duas em cada planta. Embora as populações estejam localizadas em ambientes sombreados e úmidos, as sementes precisam de luz para germinar, sendo fotoblásticas positivas. A taxa de germinação em papel de filtro na câmara de germinação foi menor (14% após 30 dias) que em solo da floresta na casa de vegetação (47% após 25 dias). Embora o fruto seja do tipo cápsula com sementes pequenas, a dispersão (anemocoria ou hidrocoria) parece não ocorrer a distâncias longas, como é observada em outras espécies de Melastomataceae com síndrome de dispersão semelhante.

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.