ABSTRACT Nectar production has an important role in pollinator attraction and successful fruit production in many self-incompatible angiosperm groups. The reproductive biology of Passiflora actinia was studied here and related to nectar dynamics. Passiflora actinia presented a temporal segregation of male and female functions at the beginning of anthesis. Due to the movements of floral verticils, the anthers were positioned in a way that favors pollination two hours before the stigmas reached the same position. The nectary consisted of an epidermis with stomata and a parenchyma rich in starch, which was hydrolyzed during anthesis. The nectary organization is probably associated with the continuous production of nectar during anthesis as well as with the high mean nectar concentration. Hand pollination tests indicated that Passiflora actinia is obligately xenogamous, depending on large bees for pollination, specifically the carpenter bee Xylocopa augusti. The continuous production of nectar may increase the number of bee visits, thus favoring pollen flow.

ABSTRACT Plant-pollinator interactions and their reproductive implications are of central importance to the organization of plant populations and communities in restoration areas. We studied the breeding system and flower visitors of Pleroma trichopodum, a pioneer species of the Atlantic Forest. We attempted to answer three questions: (1) Is P. trichopodum dependent on pollinators and mates for reproduction? (2) What are the pollinators of P. trichopodum? (3) Do tree flower-density and flowering-tree density of P. trichopodum enhance the visitation rate of focal trees and their flowers? We tested the breeding system through pollination treatments. We performed focal observations on 10 trees and analyzed the relationship between tree and flower visitation rates, and the tree flower-density and flowering-tree density with Generalized Linear Mixed Models (GLMM). Pleroma trichopodum sets fruits by selfing and outcrossing, both of which occur only through pollinator visits. Bees visited P. trichopodum, and their visitation increased with tree flower-density. Thus, individuals with greater floral displays may function as magnet species, enhancing the pollination of nearby plant species. This characteristic, allied with the ability to reproduce without a mate (selfing) and a tolerance of soggy soils, make P. trichopodum a candidate for Atlantic Forest restoration.

ABSTRACT Plant size and clonality are important traits for explaining the reproductive effort of clonal plants. Larger plants can invest more resources into reproduction, and clonality is known to increase reproductive effort. Moreover, reproductive effort is influenced by environmental variation, and so the spatial distribution of plants may affect plant reproductive effort. We investigated the effect of plant size, clonality and spatial distribution on the reproductive effort of Vriesea carinata in the Atlantic Forest in the state of Paraná, Brazil. We marked twenty individual plants and measured their rosette size, biomass and number, as well as rosette reproductive effort (number of flowers, fruits and seeds). We also evaluated the relationship between reproductive effort and spatial distribution of plants. Reproductive effort did not correlate with size, whereas greater clonal growth contributed to a lower reproductive effort because rosettes within clones that had more rosettes set fewer flowers. We found that plants growing closer to each other exhibited similar reproductive efforts independently of vegetative traits, because reproductive traits were spatially autocorrelated. In Vriesea carinata, the main drivers of reproductive effort are clonality, which decreases flower production, and spatial factors, which result in greater similarity in reproductive efforts among more proximate plants.

Resumo As florestas apresentam um enriquecimento gradual de espécies e um aumento em complexidade estrutural e funcional durante a sucessão. Este trabalho caracterizou os traços reprodutivos de plantas relacionados à polinização e à dispersão em um gradiente sucessional, em uma área de Mata Atlântica, Sul do Brasil. O levantamento das espécies foi realizado em áreas florestais sucessionais oriundas de regeneração natural, em idades variando entre 2 e 80 anos após o abandono da pastagem. Houve predominância de flores polinizadas por abelhas, abertas, hermafroditas e que disponibilizam o pólen como em todos estádios sucessionais. Os sistemas de polinização abiótico e misto, sistema sexual dióico, espécies com dispersão abiótica, com muitas sementes e sementes pequenas diminuíram com a idade das florestas. Espécies polinizadas por abelhas, flores com corola funil/sino, como de corola pequena e andróginas assim como espécies dispersas bioticamente e com sementes grandes aumentaram no gradiente sucessional. A riqueza funcional e a dispersão funcional aumentaram no gradiente sucessional. As mudanças da diversidade funcional foram relacionadas com o aumento da riqueza de espécies indicando que o enriquecimento de espécies em florestas mais velhas adicionou novos conjuntos de traços reprodutivos. Estas mudanças de distribuição de traços e de diversidade funcional no gradiente sucessional sugerem um incremento do papel de interações bióticas na estruturação de assembleias de plantas de florestas tropicais secundárias.

Abstract Niche and neutral processes shape community assembly with a possible shift of niche and neutral importance in communities undergoing temporal changes during succession. Functional diversity helps to discriminate assembly processes since trait distribution is dependent on those processes. We evaluated the changes in reproductive traits related to pollination and seed dispersal in a successional gradient in an Atlantic Forest area, Southern Brazil. We surveyed forests undergoing regeneration varying in age from 2 to 80 years after pasture abandonment. We expected an increase in functional diversity of reproductive traits and a greater role of limiting similarity across succession. Abiotic and mixed pollination systems, dioecious sexual system, biotic dispersed, many-seeded and small-seeded species decreased as the forest got older. Conversely, bee-pollinated, bell-shaped, small and androgynous flowers increased across forest succession as well biotic dispersed and large-seeded species. Functional richness and functional dispersion were higher in older forests. Changes in functional diversity were positively related to species richness, indicating that species enrichment in older forests added new sets of reproductive traits. These changes in trait distribution and functional diversity across succession in the Atlantic Forest suggest an increased role of biotic interactions and limiting similarity process structuring plant assemblages of second-growth tropical forests.

The abundance and distribution of resources for pollinators modulate the reproduction of plants. This study attempted to determine whether and how the fruiting success of Tibouchina trichopoda flowers varies due to plant size, number of flowers produced, distance between individuals and cardinal orientation of the flowers. From July 2009 to February 2010, we conducted monthly evaluations of 21 plants in an area of regenerating forest within the Atlantic Forest biome, located near the municipality of Antonina, in southern Brazil. Floral density was monitored weekly. During the study period, the species bloomed twice: once during the cool, dry season, showing an extended flowering period with scattered flowering individuals; and once during the hot, rainy season, showing a brief flowering period with a high density of flowering individuals. Plant size was not found to influence fruiting success. There was greater production of flowers and fruits during the brief flowering period. The extent of the floral display positively affected the fruiting success of the flowers, the effect being greater when the density of flowers was low and tending to stabilize at higher densities. The northern-facing portions of tree crowns produced more flowers and therefore had greater fruiting success than did the southern-facing portions. However, the proportion of fruits produced per flower was the same.

A atividade dos polinizadores é afetada pela disponibilidade de recursos. Flores que produzem mais néctar podem ser mais visitadas e assim apresentar maior produção de frutos. O efeito da produção de néctar na atividade dos polinizadores foi testado em duas espécies cultivadas de maracujá, Passiflora alata Curtis e Passiflora edulis Sims, em Morretes, Paraná. Botões foram ensacados e o néctar acumulado das flores foi coletado em intervalos de 1 h. Em P. alata o volume e a concentração de solutos no néctar aumentaram durante o período de antese, associados com o aumento da temperatura. Em P. edulis, o volume aumentou durante o período diurno da antese, e decresceu após as 18 horas. A concentração de solutos no néctar permaneceu constante. A taxa média de visitação de Xylocopa frontalis (Olivier) em P. alata foi de 1,7 visitas/100flores/hora e em P. edulis foi de 6,6 visitas/100flores/hora, sendo constante durante a antese. A taxa média de visitação de Bombus morio (Swederus) em P. alata foi de 5,8 visitas/100flores/hora, sendo mais alta no início da antese. A constância das visitas de X. frontalis deve estar associada à produção contínua de néctar em ambas as espécies de maracujazeiros. Como as espécies são xenogâmicas, a manutenção das visitas é importante para propiciar o fluxo de pólen entre indivíduos e assim garantir boa produção de frutos.

Pollinator activity is affected by resource availability. Flowers that produce more nectar are visited more, which results in a greater fruit set. The effect of nectar production on pollinator activity was tested in two cultivated species of passion fruit, Passiflora alata Curtis and Passiflora edulis Sims, in Morretes, Paraná. Flower buds were bagged and the accumulated nectar of flowers was collected hourly. The volume and concentration of nectar of P. alata increased during anthesis, which was associated with rising temperatures. The volume of nectar of P. edulis increased during the diurnal period of anthesis, falling after 6 pm. The nectar concentration was constant during anthesis. The visitation rate of Xylocopa frontalis (Olivier) was constant for P. alata (1.7 visits/100flowers/hour) and for P. edulis (6.6 visits/100flowers/hour). The visitation rate of Bombus morio (Swederus) for P. alata was 5.8 visits/100flowers/hour, and was higher at the beginning of anthesis. The constant rate of visits by X. frontalis may be associated to the continuous nectar production by both species of passionflower. Since both species are xenogamous, the maintenance of visits during anthesis may favor pollen flow among the individuals assuring good crop production.

A biologia reprodutiva, incluindo fenologia, biologia floral, polinização e sistemas reprodutivos foram estudados em Floresta com Araucária no Paraná. A fenologia e reprodução de plantas terrícolas foram acompanhadas em outubro 2006 e de maio a outubro de 2007. O pico de floração ocorreu de junho a setembro e a frutificação de junho a outubro. A antese durou um dia. As flores foram polinizadas principalmente por Stephanoxis lalandi e a borboleta mais freqüente foi Lychnuchoides ozias ozias. A concetração de nectar declinou durante a antese, enquanto que o volume de néctar permaneceu constante. Aechmea distichantha é auto-compatível com 30-45% de frutificação nos testes de autopolinização. A luz solar influenciou a reprodução: quando controlados o tamanho das bromélias e das inflorescências, plantas no sol produziram mais sementes por frutos que plantas na sombra. A reprodução também foi associada com tamanho das inflorescências quando controlado o tamanho da bromélia. Inflorescências maiores em plantas de mesmo tamanho produzem mais flores e mais sementes por fruto.

Reproductive biology, including phenology, flower biology, pollination, and the reproductive system in the bromeliad Aechmea distichantha were studied in an Araucaria forest in the state of Paraná. Phenology and reproduction in terricolous plants were followed in October 2006 and May - October of 2007. Flowering peaked from June to September and fruiting was from June to October. Flower anthesis lasted one day. Flowers were pollinated the most by the hummingbird Stephanoxis lalandi and the most common butterfly visitor was Lychnuchoides ozias ozias. Nectar concentration declined during anthesis, while nectar volume was constant. Aechmea distichantha is self-compatible with 30-45% fruit formation in self-pollination tests. Sunlight influences reproduction: when controlling for bromeliad and inflorescence size, plants in sunlight produced more seeds per fruit than plants in the shade. Reproduction was also associated with inflorescence size when controlling for bromeliad size. That is, larger inflorescences in similar sized plants produced more flowers and more seeds per fruit.

Organização da guilda de polinizadores e sua consequência para reprodução em três espécies sincropátricas de Tibouchina (Melastomataceae). Em espécies de plantas que co-florescem, a partilha de polinizadores pode resultar em transferência interespecífica de pólen e redução da fecundidade. A competição pode ser relaxada quando existe uma grande quantidade de suprimento de pólen ou se cada planta ocupa manchas distintas do hábitat. A reprodução de Tibouchina cerastifolia (Naudin) Cogn., T. clinopodifolia (DC.) Cogn. e T. gracilis (Bonpl.) Cogn. foi estudada em Floresta Atlântica para verificar se a sincronopatria levava à partilha temporal das espécies de polinizadores. Onze espécies de abelhas compuseram a guilda de polinizadores. Entre eles, houve sobreposição na composição de espécies de polinizadores e no período de visitação floral. A competição direta por pólen em Tibouchina Aubl. na área de estudo parece levar a distintos períodos de atividade entre as abelhas, onde Bombus pauloensis Friese,1913 foi mais ativa mais cedo enquanto que as outras espécies foram ativas mais tarde. Bombus pauloensis, a maior espécie de abelha registrada em nas flores de Tibouchina, foi o polinizador mais importante e eficiente. Esta espécie coletava pólen antes das outras abelhas e apresentou um menor tempo de manipulação das anteras. As plantas se reproduziram sexuadamente, por autogamia e xenogamia e a hibridação não foi inibida por reações de incompatibilidade no estilete. A ausência de competição direta por pólen e de partilha de polinizadores pode refletir uma interação de facilitação entre estas três espécies pioneiras e sincronopátricas de Tibouchina.

Pollinator guild organization and its consequences for reproduction in three synchronopatric species of Tibouchina (Melastomataceae). In co-flowering plant species, pollinator sharing can result in interspecific pollen transfer and fecundity reduction. Competition will be relaxed whenever there is a large amount of initial pollen supply or if each plant species occupies different habitat patches. Reproduction in Tibouchina cerastifolia (Naudin) Cogn., T. clinopodifolia (DC.) Cogn. and T. gracilis (Bonpl.) Cogn. was studied in an area of Atlantic rainforest to examine whether synchronopatry induces time partitioning among pollinator species. Eleven bee species comprised the pollinator guild. Among pollinators, there were overlaps in bee species composition and in flower visitation time. Direct competition for pollen in Tibouchina Aubl. at the study site seems to lead to different activity periods among the bee species, in which Bombus pauloensis Friese,1913 was most active earlier, while the other species were active later in the day. Bombus pauloensis, the largest bee species recorded on Tibouchina flowers, was the most important and efficient pollinator. This species harvested pollen before the other species and had the shortest handling time. The plants reproduced sexually by selfing or outcrossing, and hybridization was not avoided by incompatibility reactions at the style. The avoidance of direct competition for pollen and no pollinator partitioning among the synchronopatric species of Tibouchina may reflect a facilitative interaction among these pioneer plants.

Resumo O gênero Tibouchina é comum no Brasil e encontrado principalmente na Floresta Atlântica, em especial na região Sudeste. No presente estudo foi avaliada a viabilidade polínica de cinco espécies de Floresta Atlântica, T. cerastifolia, T. clinopodifolia, T. gracilis, T. pulchra e T. sellowiana, utilizando-se três soluções histoquímicas: carmim acético, Alexander e cloreto de 2,3,5-trifenil-tetrazólio. Foi avaliada a variação de viabilidade entre as anteras dimórficas, entre as soluções histoquímicas e entre os indivíduos. Em T. pulchra e T. sellowiana também foi avaliada a variação entre os dois primeiros dias a partir da abertura da flor. Nenhuma espécie apresentou variação relacionada ao dimorfismo das anteras. Nas duas espécies que apresentam maior longevidade floral, a viabilidade foi semelhante no primeiro e segundo dias em T. pulchra, enquanto que em T. sellowiana as flores de segundo dia apresentaram pólen menos viável. As soluções histoquímicas empregadas revelaram taxas de viabilidade polínica distintas. Houve variação na viabilidade polínica entre os indivíduos em todas as espécies. A solução de cloreto de 2,3,5-trifenil-tetrazólio revelou a menor taxa de viabilidade em todos os indivíduos. As soluções histoquímicas empregadas, a amostragem e a longevidade floral foram fatores relacionados à variação da viabilidade polínica.

Abstract The genus Tibouchina is common in Brazil, mainly found in Atlantic Forest, in Southeast region. In this study the pollen viability was evaluated in five species of Atlantic Forest, T. cerastifolia, T. clinopodifolia, T. gracilis, T. pulchra and T. sellowiana, using three histochemical solutions: acetocarmine, Alexander and 2, 3, 5-triphenyltetrazolium chloride. The variation of pollen viability was evaluated among the dimorphics anthers and among individuals. In T. pulchra and T. sellowiana it was also evaluated the variation among the two first days after flower openning. There was no variation related to anther dimorphism. Considering the two species with the longest flower longevity, the viability was similar in the first and second day in T. pulchra, but in T. sellowiana the pollen from the second day was less viable. The histochemical tests employed showed distinct rates of pollen viability. Rates of pollen viability were distinct among individuals in each species. The stain 2, 3, 5-triphenyltetrazolium chloride showed the smaller viability. The results show that the treatments employed, sampling and flower longevity were factors related to pollen viability.

Neste trabalho foram investigados os sistemas reprodutivos de espécies de Melastomataceae ocorrentes em formações florestais da Serra do Japi. Esta investigação foi baseada em polinizações controladas e em análises de viabilidade de pólen e de crescimento de tubo polínico. Entre as populações de 13 espécies estudadas, sete mostraram produção de frutos apomíticos (Leandra australis, L. melastomoides, L. purpurascens, Miconia latecrenata, M. petropolitana, Ossaea amygdaloides e O. confertiflora). Entre as seis espécies não apomíticas, apenas uma apresentou mecanismo de auto-incompatibilidade (Miconia pusilliflora), enquanto que as demais se mostraram auto-compatíveis (Leandra dasytricha, L. regnellii, Tibouchina cerastifolia, T. sellowiana e T. semidecandra). A análise de pólen mostrou que as espécies apomíticas produzem uma proporção de grãos viáveis significativamente menor do que aquelas que não são apomíticas.

The breeding systems of some Melastomataceae species were investigated in populations from forests on the Serra do Japi, southeastern Brazil. This study was based on controlled pollinations and analysis of pollen viability and pollen-tube growth. Among populations of the 13 species studied, seven showed apomictic fruits (Leandra australis, L. melastomoides, L. purpurascens, Miconia latecrenata, M. petropolitana, Ossaea amygdaloides e O. confertiflora). Among the six non-apomictic species only one showed self-incompatibility (Miconia pusilliflora), whereas the others showed self-compatibility (Leandra dasytricha, L. regnellii, Tibouchina cerastifolia, T. sellowiana e T. semidecandra). Pollen analysis showed that the apomictic species have significantly lower viability levels than the non-apomictic species.

RESUMO A polinização de Passiflora alata Dryander decorre da atividade de coleta de néctar ou pólen por abelhas Anthophoridae e Apidae de grande porte durante todo o período de floração entre março e setembro. O vôo, possivelmente seguindo rotas de visitas, permite a polinização cruzada, favorecida pela deflexão dos órgãos reprodutivos. Há flores que não apresentam deflexão dos estiletes e podem ser consideradas como funcionalmente masculinas, o que pode interferir na alocação total de recursos para a produção de sementes.

ABSTRACT The pollination of Passiflora alata Dryander is accomplished by Anthophoridae and Apidae large bees when collecting nectar or pollen during the flowering time from march to setember. Forage fligth behavior of trap-line bees possibly allowded cross-pollination what is enhanced by deflection of the reproductive organs. Styles of some flowers may not be deflected and could be considerd as functionally male ones, what may interfer in resource allocation to seedset.