Phenology, floral and reproductive biology of Dichorisandra rhizantha Aona (Commelinaceae), an endemic endangered species in an Atlantic Forest fragment.

Commelinaceae is an important component of the Atlantic Forest and its analysis can provide information on the conservation of that species. This study had the objective of analyzing the phenology and floral and reproductive biology of Dichorisandra rhizantha, growing in an Atlantic Forest fragment, to obtain data related to the form of reproduction and pollination mechanism of this species. We monitored the phenophases and reproductive biology of the D. rhizantha in the area studied using different methodologies and microscopy techniques (LM, SEM and FM). The flowering period occurs during the dry season. The species is andromonoecious and has daytime anthesis (4:30 a.m.-1:30 p.m.). It has purple zygomorphic flowers and rimose but functionally poricidal anthers. The pollen viability was 97.6% and the two floral morphs investigated contained a large quantity of pollen grains. The time of greatest stigma receptivity (anthesis) coincided with the pollen viability. Cross-pollination was the only efficient reproductive system of the species, with observation of gametophytic self-incompatibility and pollen tubes growth only as far as the middle or lower third of the pistil. We observed bee species of the genera Xylocopa and Euglossa visiting the flowers and acting as effective pollinators. The immature fruits were formed after about 3.25 ± 1 days, and the presence was noted of ants of the genus Cephalotes acting as dispersers of the seeds, with aril serving as the reward. Although the natural reproductive system is efficient, D. rhizantha is endangered, mainly due to fragmentation of habitat caused by deforestation and urban encroachment.

Hummingbird contribution to plant reproduction in the rupestrian grasslands is not defined by pollination syndrome.

Floral traits mediate the roles of distinct animals as effective pollinators along a generalization/specialization continuum. Many plant species are visited by different pollinator functional groups and the specific contribution of each group is expected to reflect the set of floral characteristics defined by pollination syndromes. Although considered a highly specialized nectarivorous group, hummingbirds frequently visit flowers lacking apparent specialization to bird pollination. How they contribute to the reproduction of these plants, however, has not been evaluated through field experiments considering multiple non-related plant species simultaneously. Here, we investigated hummingbirds' contributions to the pollination of ten plant species comprising a gradient of adaptation to bird pollination in the Brazilian rupestrian grasslands. We excluded hummingbirds from flowers and evaluated their relative contribution in comparison to insects (mainly bees) on conspecific/heterospecific pollen deposition and fruit set. Floral traits that are typically associated with bird pollination were associated with increased pollen deposition, but not with fruit set in the presence of hummingbirds. With hummingbirds, conspecific and heterospecific pollen deposition increased in most species, while fruit set increased in four plant species with varying degrees of fit to ornithophily. Our results show that assessing the relative contribution of specific pollinator groups may depend on when this contribution is measured, i.e. pollen deposition or fruit set. Considering fruit set, our results indicate that hummingbirds contributed to plant reproduction independently of the fit to bird pollination syndrome. This emphasizes their importance as under-appreciated generalized pollinators in some communities.

Ant pollination of Paepalanthus lundii (Eriocaulaceae) in Brazilian savanna.

BACKGROUND AND AIMS: Ant-plant associations are widely diverse and distributed throughout the world, leading to complex ecological networks. Regarding ant-plant mutualism, ant pollination is a very rare interaction and few studies have shown the role of ants as pollinators. Therefore, we aimed to evaluate the role of ants as effective pollinators of Paepalanthus lundii (Eriocaulaceae) in a Brazilian savanna. METHODS: Fieldwork with experimental manipulation was conducted to evaluate the fitness of P. lundii, considering potential pollinators. For this, we mainly observed the number of seeds produced in different conditions: control, ant exclusion, exclusion of flying insects, and exclusion (entomophily test) of both ants and flying insects. Furthermore, we evaluated all floral visitors throughout the day, stigma receptivity, the numbers of male and female flowers, and patterns of species co-occurrence, which can indicate the presence of different pollinators in the plants at the same time. KEY RESULTS: We observed a relation between seed production and ant visits; Camponotus crassus was the most frequent floral visitor and the most effective pollinator. Also, we observed a statistical difference between the numbers of male and female flowers produced, with a greater number of male flowers. Furthermore, P. lundii presented flowering asynchrony, with 12 different types of maturation sequence, which indicates a cross-pollination system. Lastly, we observed an overlap of the greatest abundance of C. crassus and the time of plant stigmatic receptivity, and a pattern of non co-occurrence of ants, which shows the pollinator role of this ant. CONCLUSIONS: Our data provide evidence that previous generalizations neglecting the importance of ants as pollinators are wrong. Brazilian savanna can reveal a lot about the ant-pollination syndrome, since this environment presents peculiar characteristics related to this association. Thus, this study has great significance for the understanding of the ant-pollination syndrome, and for the understanding of the complex ecological networks present in these dry arid systems.

Assessing the geographic dichotomy hypothesis with cacti in South America.

The Cactaceae is one of the most conspicuous and ecologically important plant families in the world. Its species may have specialist or generalist pollination systems that show geographic patterns, which are synthesised in the Geographic Dichotomy Hypothesis. Here, we assess this hypothesis in five countries in both tropical and extratropical regions, evaluating the pollinator visitation rate and pollinator identity and abundance. We calculate the Shannon diversity index (H') and evenness (J) and evaluate differences between latitude parameters with a Student t-test. Overall, we found more specialised pollination systems in all tropical sites; the richness, diversity and evenness of pollinators was reduced in comparison to extratropical regions, where the pollination system was generalised. Our results support the geographic dichotomy hypothesis in the cacti of South America, suggesting that environmental factors underlying the latitudinal patterns can help to explain differences in the pollination syndrome between tropical and extratropical regions.

Reproductive biology and nectar secretion dynamics of Penstemon gentianoides (Plantaginaceae): a perennial herb with a mixed pollination system?

BACKGROUND: In many plant species, pollination syndromes predict the most effective pollinator. However, other floral visitors may also offer effective pollination services and promote mixed pollination systems. Several species of the species-rich Penstemon (Plantaginaceae) exhibit a suite of floral traits that suggest adaptation for pollination by both hymenopterans and hummingbirds. Transitions from the ancestral hymenopteran pollination syndrome to more derived hummingbird pollination syndrome may be promoted if the quantity or quality of visits by hummingbirds is increased and if the ancestral pollinator group performs less efficiently. The quantification of such shifts in pollination systems in the group is still limited. We aimed to investigate floral traits linked to this pollination syndrome in Penstemon gentianoides with flowers visited by bumblebees and hummingbirds. METHODS: We investigated the floral biology, pollinator assemblages, breeding system and nectar production patterns of P. gentianoides inhabiting a temperate montane forest in central Mexico. Pollination experiments were also conducted to assess the pollinator effectiveness of bumblebees and hummingbirds. RESULTS: P. gentianoides flowers are protandrous, with 8-d male phase (staminate) flowers, followed by the ∼1-7 d female phase (pistillate phase). Flowers display traits associated with hymenopteran pollination, including purple flowers abruptly ampliate-ventricose to a broad throat with anthers and stigmas included, and long lifespans. However, the nectar available in the morning hours was abundant and dilute, traits linked to flowers with a hummingbird pollination syndrome. Two hummingbird species made most of the visits to flowers, Selasphorus platycercus (30.3% of all visits), followed by Archilochus colubris (11.3%). Bumblebees (Bombus ephippiatus, B. huntii and B. weisi) accounted for 51.8% of all recorded visits, but their foraging activity was restricted to the warmer hours. Hummingbirds made more foraging bouts and visited more flowers than hymenopteran species. Flowers experimentally pollinated by B. ephippiatus produced significantly more fruits than those pollinated by S. platycercus. However, there was no statistical difference in the number of seeds produced per fruit when a bumblebee or a hummingbird was the pollinator. CONCLUSIONS: We have shown that bumblebees and hummingbirds visit and pollinate P. gentianoides flowers. Despite floral traits resembling the hymenoptera pollination syndrome, flowers of P. gentianoides offer characteristic nectar rewards to flowers with a hummingbird pollination syndrome. Although pollination efficiency is higher among flowers visited by hymenoptera, the noteworthy percentage of fruit production and number of seeds per fruit derived from hummingbird pollination highlights the importance of hummingbirds as a functional group of pollinators that might have potential evolutionary consequences to the plants.

POLINIZACIÓN POR ABEJAS EN Syagrus orinocensis (ARECACEAE) EN LA ORINOQUIA COLOMBIANA / Bee pollination in Syagrus orinocensis (Arecaceae) in the Colombian Orinoquia

RESUMEN La ecología de la polinización de Syagrus orinocensis fue estudiada durante tres períodos reproductivos, en un bosque de piedemonte en Casanare, Orinoquia colombiana. Syagrus orinocensis crece hasta 10m., produce de una a cuatro inflorescencias bisexuales aunque ocasionalmente son unisexuales. Las bisexuales presentan flores estaminadas y pistiladas agrupadas en tríadas, y las unisexuales llevan sólo estaminadas en díadas. Las inflorescencias protándricas abren de día, y permanecieron activas 26 días; la fase masculina se extendió los primeros 15 días, siguió una fase inactiva por ocho días, y finalmente, la fase pistilada por tres días. Las inflorescencias fueron visitadas por 43 especies de insectos de los órdenes Coleoptera, Hymenoptera y Diptera. Los insectos visitaron las inflorescencias en fase masculina, donde buscaban polen y lugares para reproducirse, mientras que en la fase femenina buscaban néctar. Los polinizadores más efectivos fueron abejas meliponinas (Apidae, Meliponini), que transfirieron adherido a su cuerpo el 83 % del polen. La presencia, constancia y eficiencia de abejas durante el estudio son evidencia sólida de melitofilia en palmas. Este caso de melitofilia permitió proponer los criterios para redefinir este tipo de polinización en palmas silvestres neotropicales.

ABSTRACT The pollination ecology of the Syagrus orinocensis was studied in the course of three consecutive yearly flowering seasons in a foothill forest in Casanare, Colombian Orinoco region. Syagrus orinocensis palms grow up to 10 m high and produce one to four bisexual, occasionally unisexual, inflorescences. The bisexual inflorescences bear staminate and pistillate flowers arranged in triads, whereas the unisexual inflorescences carry only staminate flowers in dyads. The inflorescences are protandric and open during daytime, remaining active for 26 days. The male phase extends for the first 15 days, which are followed by 8 days of an inactive phase; the pistillate phase lasts up to three days. The inflorescences of S. orinocensis were visited by 43 species of insects belonging to the orders Coleoptera, Hymenoptera and Diptera. The presence of anthophilous insects was primarily restricted to the male phase of anthesis, during which the visitors searched for pollen and breeding sites; those which visited inflorescences during the female phase seeked out nectar. The most effective pollinators of S. orinocensis were stingless bees (Apidae, Meliponini), as they transferred in average 83% of the pollen that reached receptive inflorescences. The presence, constancy and efficiency of stingless bees during this study constitute solid evidence of melittophily in S. orinocensis and allows us to propose criteria to redefine this pollination syndrome in Neotropical wild palms.

Pollination and dispersal systems in a Cerrado remnant (Brazilian Savanna) in Southeastern Brazil

The aim of this work was to identify the pollination and dispersal systems among the species in a disjunct marginal cerrado area and to compare the frequency of these systems to those found in other Neotropical vegetation. The floral and diaspore traits and the pollination and dispersal systems of 176 species were analysed in a cerrado remnant in Southeastern Brazil. The most frequent pollination system was melittophily (63 percent of the studied species) with the remaining 37 percent distributed among diverse pollination systems. Zoochory was the predominant system of dispersal (44.9 percent). The frequencies of melittophily and zoochory observed in diverse tropical areas were the main feature that allowed the formation of distinct groups in the dendrograms generated by cluster analysis.