Nectar-mediated avian pollination in Cattleya (Orchidaceae: Laeliinae).

Orchids are notable for the variety of rewards produced for their pollinators. Nevertheless, many species are rewardless and pollinated by food-deceptive strategies. This is the case for Cattleya, a Neotropical genus of more than 100 species supposedly pollinated by food-deception. Here, I studied a member of the clade Sophronitis (Cattleya cernua) in forest areas of southeastern Brazil. The study included analyses of floral morphology, light microscopy of secretory tissues, histochemistry of flower resources, and measurements of nectar volume and concentration. Data on pollinators and mechanisms of pollination were collected in the field by means of film records, while breeding systems were studied using experimental treatments applied to cultivated specimens. The flower traits of Cattleya cernua show strong adaptation to avian pollination. Flowers produce 1-7 µl of a moderately concentrated nectar (17-50%). A secretory tissue at the distal portion of the cuniculus is responsible for nectar secretion. Secreted nectar is stored in the nectar chamber. Pollinaria deposition on the apex of bird beaks allows both short- and long-billed birds to pollinate the flowers. Plants from the studied population were fully self-compatible but pollinator-dependent. Pollinator frequency was higher than in food-deceptive Cattleya. As far as is known, this is the first record of nectar production in Cattleya. The discovery of a reward-producing species (C. cernua) within a genus entirely pollinated by food-deception strategies (Cattleya) provides clues and new insights into understanding the diversification of Laeliinae across the Neotropics.

Linking pollen limitation and seed dispersal effectiveness.

Seed production and dispersal are crucial ecological processes impacting plant demography, species distributions and community assembly. Plant-animal interactions commonly mediate both seed production and seed dispersal, but current research often examines pollination and seed dispersal separately, which hinders our understanding of how pollination services affect downstream dispersal services. To fill this gap, we propose a conceptual framework exploring how pollen limitation can impact the effectiveness of seed dispersal for endozoochorous and myrmecochorous plant species. We summarize the quantitative and qualitative effects of pollen limitation on plant reproduction and use Optimal Foraging Theory to predict its impact on the foraging behaviour of seed dispersers. In doing so, we offer a new framework that poses numerous hypotheses and empirical tests to investigate links between pollen limitation and seed dispersal effectiveness and, consequently, post-dispersal ecological processes occurring at different levels of biological organization. Finally, considering the importance of pollination and seed dispersal outcomes to plant eco-evolutionary dynamics, we discussed the implications of our framework for future studies exploring the demographic and evolutionary impacts of pollen limitation for animal-dispersed plants.

Non-species-specific pollen transfer and double-reward production in euglossine-pollinated Vanilla.

Commonly attributed to orchids, the pollen movement in Vanilla has been associated with food deception and specific plant-pollinator relationships. This study investigated the role of flower rewards and pollinator specificity in the pollen transfer of a widely distributed member to the euglossinophilous Vanilla clade, V. pompona Schiede using data collected from Brazilian populations. These included investigations on morphology, light microscopy and histochemistry, and analysis of flowers scent using GC-MS. The pollinators and the mechanisms of pollination were recorded through focal observations. The yellow flowers of V. pompona are fragrant and offer nectar as reward. The major volatile compound of the V. pompona scent, carvone oxide, shows convergent evolution in Eulaema-pollinated Angiosperms. The pollination system of V. pompona is not species-specific, but its flowers are strongly adapted to pollination by large Eulaema males. Pollination mechanism is based in a combination of perfume collection and nectar seeking. The dogma of a species-specific pollination system based on food deception in Vanilla has been broken with the increase in studies on this Pantropical orchid genus. Here, least three bee species and dual reward-offering are involved in pollen transfer in V. pompona. Visitation frequency of bees collecting perfumes, used in courtship by male euglossines, is higher than in searching for food, as short-lived young euglossine males seem to be more interested in sex than food. A pollination system based on offering both nectar and perfumes as resources is described for the first time in orchids.

Comparative reproductive biology and pollinator specificity among sympatric Gomesa (Orchidaceae: Oncidiinae).

The incredible pollination mechanisms displayed by orchid flowers has inspired biologists over the centuries. Based on the intriguing flower structures, the relationship among orchid species and their pollinators has been frequently regarded as very specialised. Given that visits on flowers pollinated by oil-collecting bees are regularly rare, and in Oncidiinae the flowers frequently attractexclusively species that act as effective pollinators, the comparative reproductive biology and pollinator specificity of two sympatric Gomesa (G. varicosa and G. montana; Oncidiinae) were analysedbased on records of floral morphology, production of floral rewards, pollinators and pollination mechanisms. Furthermore, experimental pollinations were carried out in order to examine the breeding systems. The results have show that in the studied population, both Gomesa are visited by several bee species, but these orchids present a specific pollination system.Pollinaria are deposited on the head of Centridini (G. varicosa and G. montana) and Epicharitini (G. varicosa) bees when landed on the central callus of the labellumto collect lipoidal substances produced by glandular elaiophores on lateral lobes of the labellum. Both species are dependent on a biotic pollen vector to set fruits. Gomesamontana is completely self-incompatible, while G. varicosa is partially self-compatible. Our results indicate that although the occurrence of self-sterile species seems to be common in Oncidiinae, in partially self-incompatible species, as is the case of G. varicosa, self-compatibility has been considered as an important factor favouring reproductive assurance in populations with low visitation frequencies, despite occurrence of inbreeding depression.

Floral biology of two Vanilloideae (Orchidaceae) primarily adapted to pollination by euglossine bees.

Vanilloideae comprises 15 genera distributed worldwide, among which are Vanilla and Epistephium (tribe Vanilleae). Based on field and laboratory investigations, the pollination biology of V. dubia and E. sclerophyllum was analysed. The former was surveyed in a semi-deciduous mesophytic forest at the biological reserve of Serra do Japi and in a marshy forest at the city of Pradópolis, southeastern Brazil. The latter was examined in rocky outcrop vegetation in the Chapada Diamantina, northeastern Brazil. In the studied populations, the tubular flowers of V. dubia and E. sclerophyllum were pollinated by bees. Pollen was deposited on either their scutellum (V. dubia) or scutum (E. sclerophyllum). The mentum region of V. dubia is dry, whereas that of E. sclerophyllum presents a small quantity of dilute nectar. Flowers of E. sclerophyllum are scentless, while those of V. dubia are odoriferous. Although V. dubia is self-compatible, it needs a pollinator to produce fruit. In contrast, E. sclerophyllum sets fruit through spontaneous self-pollination, but biotic pollination also occurs. Both species are primarily adapted to pollination by euglossine bees. Pollination by Euglossina seems to have occurred at least twice during the evolution of Vanilleae. Furthermore, shifts between rewarding and reward-free flowers and between autogamous and allogamous species have been reported among vanillas.

Community-wide assessment of pollen limitation in hummingbird-pollinated plants of a tropical montane rain forest.

BACKGROUND AND AIMS: Although pollen limitation of reproduction (PL) has been widely studied, our understanding of its occurrence in tropical communities, especially for bird-pollinated plants, is underdeveloped. In addition, inclusion of both quantity and quality aspects in studies of PL are generally lacking. Within hummingbird-pollinated plants, a prediction was made for higher PL for the quality than quantity aspects and a minor effect of temporal variation because hummingbirds are constant and efficient pollen vectors but they may transfer low quality pollen. METHODS: Field hand and open pollination experiments were conducted on 21 species in a tropical montane rain forest over 2 years. The quantity (fruit set and seeds per fruit) and quality (seed weight and germination) aspects of reproduction were assessed as the response to open pollination relative to outcross hand pollination. The relationships between the effect size of quantity and quality aspects of reproduction and predictive plant features (self-incompatibility, autogamy, density and pollinator specialization level) were assessed with phylogenetic generalized linear models. KEY RESULTS: Just over half of all the species expressed PL for one or more response variables. On average, the severity of PL was strong for one quality variable (seed germination; 0·83), but insignificant for another (seed weight; -0·03), and low to moderate for quantity variables (0·31 for seeds per fruit and 0·39 for fruit set). There was only a minor contribution of temporal variation to PL within the studied species. Common predictors of PL, i.e. phylogenetic relatedness, self-incompatibility, autogamy, plant density and pollinator specialization level, did not adequately explain variation in PL within this community. CONCLUSIONS: Despite the measurable degree of PL within these hummingbird-pollinated plants, the causes of pollen quality and quantity insufficiency are not clear. Variables other than those tested may contribute to PL or causes of PL may vary among species and cannot adequately be accounted for when assessed from the within-community perspective.

Pollination and breeding system of Couepia uiti (Mart. and Zucc.) Benth (Chrysobalanaceae) in the Pantanal da Nhecolândia

The pollination biology and breeding system of Couepia uiti was studied. In this species, flowers opened at 06:00 AM anthesis, and nectar production began at around 0800 h, reached a maximum volume from 09:30 AM to 10:30 AM, and decreased thereafter. The nectar sugar concentration increased continuously, but showed an abrupt increase from 10:00 AM to 12:00 AM. Pollen release occurred at about 09:30 AM and was quickly collected. The stigmas became receptive at around 12:00 AM. The pollinators of C. uiti included the bees Apis mellifera, Xylocopa sp. and Bombus sp., and three species of wasps. This conclusion was based on the observation that these hymenopterans had C. uiti pollen on their bodies, visited the receptive flowers, and touched the anthers and stigmas, thereby promoting pollination. Of these floral visitors, A. mellifera was considered to be the most efficient pollinator. However, mixed pollination also occurred. The number of C. uiti flowers visited in the morning (n = 52) was three times smaller than in the afternoon (n = 62), and the species richness of floral visitors was also bigger in the afternoon (eight in the afternoon versus five in the morning). This finding indicated that these floral visitors preferred to exploit nectar rather than pollen. Controlled pollination experiments showed that C. uiti was a self-incompatible species that produced fruits only by cross-pollination. Treatments such as agamospermy and spontaneous and self-pollinations did not produce fruits.

Estudei a biologia da polinização e o sistema reprodutivo de Couepia uiti. Esta espécie apresentou antese matutina, produção de néctar iniciou por volta das 08:00 horas, atingindo volume máximo entre 09:30 e 10:30 horas e diminuindo após este horário. A concentração de açúcares no néctar aumentou continuamente, apresentando aumento abrupto entre 10:00 e 12:00 horas. A liberação de pólen ocorreu às 09:30 horas e este foi rapidamente removido. O estigma estava receptivo às 12:00 horas. Considerei polinizadores de C. uiti as abelhas: Apis mellifera, Xylocopa sp., Bombus sp. e três espécies de vespas, pois apresentaram polén de C. uiti em seus corpos. Visitaram flores quando estavam receptivas, tocando as anteras e os estigmas, promovendo a polinização. Dentre os visitantes florais, A. mellifera foi considerado o polinizador mais eficiente, no entanto, polinização mista parece ocorrer. O número de visitas a flores de C. uiti pela manhã (n = 52) foi três vezes menor que o encontrado no período da tarde (n = 162) e o número de espécies de visitantes também foi maior à tarde, sendo encontradas cinco espécies durante a manhã e oito à tarde. Portanto, é possível concluir que estes visitantes florais preferem explorar o néctar em comparação ao pólen. Os tratamentos de polinizações controladas mostraram que C. uiti é uma espécie auto-incompatível, produzindo frutos somente por polinização cruzada. Tratamentos de agamospermia, polinização espontânea e autopolinização não produziram frutos.

Pollination and breeding system of Couepia uiti (Mart. and Zucc.) Benth (Chrysobalanaceae) in the Pantanal da Nhecolândia

The pollination biology and breeding system of Couepia uiti was studied. In this species, flowers opened at 06:00 AM anthesis, and nectar production began at around 0800 h, reached a maximum volume from 09:30 AM to 10:30 AM, and decreased thereafter. The nectar sugar concentration increased continuously, but showed an abrupt increase from 10:00 AM to 12:00 AM. Pollen release occurred at about 09:30 AM and was quickly collected. The stigmas became receptive at around 12:00 AM. The pollinators of C. uiti included the bees Apis mellifera, Xylocopa sp. and Bombus sp., and three species of wasps. This conclusion was based on the observation that these hymenopterans had C. uiti pollen on their bodies, visited the receptive flowers, and touched the anthers and stigmas, thereby promoting pollination. Of these floral visitors, A. mellifera was considered to be the most efficient pollinator. However, mixed pollination also occurred. The number of C. uiti flowers visited in the morning (n = 52) was three times smaller than in the afternoon (n = 62), and the species richness of floral visitors was also bigger in the afternoon (eight in the afternoon versus five in the morning). This finding indicated that these floral visitors preferred to exploit nectar rather than pollen. Controlled pollination experiments showed that C. uiti was a self-incompatible species that produced fruits only by cross-pollination. Treatments such as agamospermy and spontaneous and self-pollinations did not produce fruits.

Estudei a biologia da polinização e o sistema reprodutivo de Couepia uiti. Esta espécie apresentou antese matutina, produção de néctar iniciou por volta das 08:00 horas, atingindo volume máximo entre 09:30 e 10:30 horas e diminuindo após este horário. A concentração de açúcares no néctar aumentou continuamente, apresentando aumento abrupto entre 10:00 e 12:00 horas. A liberação de pólen ocorreu às 09:30 horas e este foi rapidamente removido. O estigma estava receptivo às 12:00 horas. Considerei polinizadores de C. uiti as abelhas: Apis mellifera, Xylocopa sp., Bombus sp. e três espécies de vespas, pois apresentaram polén de C. uiti em seus corpos. Visitaram flores quando estavam receptivas, tocando as anteras e os estigmas, promovendo a polinização. Dentre os visitantes florais, A. mellifera foi considerado o polinizador mais eficiente, no entanto, polinização mista parece ocorrer. O número de visitas a flores de C. uiti pela manhã (n = 52) foi três vezes menor que o encontrado no período da tarde (n = 162) e o número de espécies de visitantes também foi maior à tarde, sendo encontradas cinco espécies durante a manhã e oito à tarde. Portanto, é possível concluir que estes visitantes florais preferem explorar o néctar em comparação ao pólen. Os tratamentos de polinizações controladas mostraram que C. uiti é uma espécie auto-incompatível, produzindo frutos somente por polinização cruzada. Tratamentos de agamospermia, polinização espontânea e autopolinização não produziram frutos.

Reproductive biology in species of Bidens L. (Asteraceae)

Studies about reproductive biology of weed species can have implications on the establishment of controlling practices that minimize the effects of these weed populations on agricultural fields. The pollination biology of Bidens alba (L.) DC., B. pilosa L., and Bidens subalternans DC., was studied at different sites and climatic seasons. Bidens pilosa and B. subalternans are widely distributed in agricultural areas, in disturbed habitats, and along road sides. Bidens alba occur only along the coast. The three species are self-compatible and non agamospermous. The composition of the pollinator community changes during the year and between sites. Hymenopterans and lepidopterans are the most frequent visitors to Bidens species in both areas studied. Although the species are self-compatible, the presence of pollinators may affect the levels of inbreeding. The attraction of insects by Bidens species may be benefical to agricultural crop and may also have important implications for conservation biology.

Estudos de biologia reprodutiva de espécies invasoras podem ter implicações sobre o estabelecimento de práticas de controle que minimizem o efeito das populações destas espécies em áreas agrícolas. A biologia da polinização de Bidens alba (L.) DC., B. pilosa L. e Bidens subalternans DC. foi estudada em diferentes locais e estações climáticas. Bidens pilosa e B. subalternans são espécies amplamente distribuídas em áreas agrícolas, em habitats perturbados e em margens de estradas. Bidens alba ocorre somente em regiões litorâneas. As três espécies são auto-compatíveis e não são agamospérmicas. A composição da comunidade de polinizadores apresenta diferenças durante o ano e entre locais. Himenópteros e lepidópteros são os visitantes mais freqüentes nas espécies de Bidens. Embora as espécies sejam auto compatíveis, a presença de polinizadores pode afetar os níveis de endocruzamento. A atração de insetos por espécies de Bidens pode beneficiar as culturas agrícolas e apresentar importantes implicações para a biologia da conservação.

Reproductive biology in species of Bidens L. (Asteraceae)

Studies about reproductive biology of weed species can have implications on the establishment of controlling practices that minimize the effects of these weed populations on agricultural fields. The pollination biology of Bidens alba (L.) DC., B. pilosa L., and Bidens subalternans DC., was studied at different sites and climatic seasons. Bidens pilosa and B. subalternans are widely distributed in agricultural areas, in disturbed habitats, and along road sides. Bidens alba occur only along the coast. The three species are self-compatible and non agamospermous. The composition of the pollinator community changes during the year and between sites. Hymenopterans and lepidopterans are the most frequent visitors to Bidens species in both areas studied. Although the species are self-compatible, the presence of pollinators may affect the levels of inbreeding. The attraction of insects by Bidens species may be benefical to agricultural crop and may also have important implications for conservation biology.

Estudos de biologia reprodutiva de espécies invasoras podem ter implicações sobre o estabelecimento de práticas de controle que minimizem o efeito das populações destas espécies em áreas agrícolas. A biologia da polinização de Bidens alba (L.) DC., B. pilosa L. e Bidens subalternans DC. foi estudada em diferentes locais e estações climáticas. Bidens pilosa e B. subalternans são espécies amplamente distribuídas em áreas agrícolas, em habitats perturbados e em margens de estradas. Bidens alba ocorre somente em regiões litorâneas. As três espécies são auto-compatíveis e não são agamospérmicas. A composição da comunidade de polinizadores apresenta diferenças durante o ano e entre locais. Himenópteros e lepidópteros são os visitantes mais freqüentes nas espécies de Bidens. Embora as espécies sejam auto compatíveis, a presença de polinizadores pode afetar os níveis de endocruzamento. A atração de insetos por espécies de Bidens pode beneficiar as culturas agrícolas e apresentar importantes implicações para a biologia da conservação.