New insights into the functional morphology of the lever mechanism of Salvia pratensis (Lamiaceae)

BACKGROUND AND AIMS: The functional morphology of Salvia pratensis flowers was re-investigated, after new insights revealed that pollen dispensing is one of the main functions of the staminal lever. In particular, no detailed information was available regarding the process of pollen transfer and the forces arising between the pollen-bearing thecae and the pollinating bee's body. The assumption was made that these forces play a significant role in pollen dispensing. METHODS: The functional morphology of S. pratensis flowers and the interaction between flowers and bees (Apis mellifera) were studied by reconstructing stress and strains by using qualitative and semi-quantitative theoretical analysis. Flowers were manipulated to study the spatial arrangement of the filament and lever, and of the head and proboscis of the visiting bee inside the tube. Photographs and films of bee visits on flowers were used to analyse the interaction of pollinator and staminal lever. KEY RESULTS: The spoon-shaped lower lever of S. pratensis has a small hole through which a bee introduces its proboscis into the corolla tube. Although mentioned for the first time by Kerner von Marilaun in 1891, presented here is the first drawing and the first photograph showing this interaction in detail. The analysis of the interaction of flower visitor and the lever mechanism revealed that the position of bees on different flowers is spatially very similar. Flower morphology constrains postures of legitimately nectar-probing bees within narrow bounds. A theoretical discussion on structural elements and force progression in the flower allows the principles of lightweight architecture in flower morphology to be recognized. CONCLUSIONS: The staminal lever of S. pratensis is a pollen-dispensing device. It seems to influence the amount of pollen deposited on pollinators by determining the forces arising between the pollinator and the pollen. The relevant forces occur either during the first, dynamic phase or during the second, almost static phase of a flower visit.     

The staminal lever mechanism in Salvia L. (Lamiaceae): a key innovation for adaptive radiation?

Floral key innovations play a significant role in the discussion of adaptive radiation in plants. The paper brings together a brief review of morphological key innovations in plants, elucidating their evolutionary significance in flower–pollinator interactions, and new data on Salvia, a genus being examined as an example for presumed adaptive radiation. We hypothesize that the characteristic staminal lever mechanism functions as a key innovation. It is defined as a functional unit including the modification of stamens to lever-like structures, their reversible movement, and the organization of the remaining floral structures involved in the process of pollen transfer. We follow the assumption that structure and functioning of the staminal levers play a major role in the process of pollen deposition on the pollinator's body, and that minute changes of both their proportions and their interactions with pollinators may have significant consequences for the pollination system. The functioning of the staminal lever mechanism is tested by field investigations, biomechanical experiments and pollination simulations. First results are presented, and possible modes of allopatric and sympatric speciation are discussed, based on morphometry of Salvia flowers and pollinators as well as on the operating mode of the staminal lever mechanism. Special attention is given to species-specific patterns of pollen deposition on the pollinator's body. We assume that, depending on the precision of the lever movement, sympatric Salvia species flowering during overlapping periods and sharing the same pollinating species may be either mechanically isolated from each other or able to hybridize. The latter may result in speciation, as may spontaneous mutations influencing the flower-pollinator interaction, e.g. by significant changes in morphometry of the staminal lever system and/or other flower structures. As a consequence, Salvia individuals may deposit pollen on a different part of the pollinator's body, or even adapt to a new pollinator species, both resulting in reproductive isolation from the parental population.     

Is pollen removal or seed set favoured by flower longevity in a hummingbird-pollinated Salvia species?

Background and Aims

The period between the beginning of anthesis and flower senescence modulates the transport of pollen by pollinators among conspecific flowers, and its length may therefore influence reproductive success. This study evaluated whether floral longevity favours pollen removal from the anthers over fecundity (seed set) in an ornithophilous species that does not undergo pollen limitation.

Methods

Field investigations were conducted on floral longevity, nectar production, pollinator behaviour, and variations in fruit set (FS), mean number of seeds per fruit (MSF) and pollen removal by hummingbirds (PR) during the anthesis of Salvia sellowiana in south-east Brazil.

Key Results

Anthesis of flowers exposed to pollinators lasted 4 d, as well as on flowers with pollen removed from the anthers or deposited on the stigma. The longevity of bagged flowers was significantly higher (approx. 9 d). FS and PR reached 87·2 and 90 %, respectively, in natural conditions. PR increased gradually over the period of anthesis; however, FS and MSF reached their maxima in the first hours of anthesis. Nectar production was continuous, but the secretion rate was reduced after pollination. The removal of nectar from non-pollinated flowers stimulated its production.

Conclusions

The longevity of anthesis in S. sellowiana seems to be related to the mechanism of gradual dispensing of pollen, resulting in greater male reproductive success. This is in agreement with the pollen-donation hypothesis. The small number of ovules (four) of S. sellowiana and the high frequency and the foraging mode of its pollinators may favour the selection for floral longevity driven by male fitness in this system.     

鼠尾草属不同物种的雄蕊杠杆机制对传粉者空间变异的进化响应

被子植物虫媒传粉植物的物种分化通常被认为是花性状响应传粉环境(传粉者)的空间变异而发生适应性分化的结果。通过对鼠尾草属(Salvia) 3个物种(共4个居群)传粉互作系统的比较, 探索了花性状对不同传粉环境的进化响应。结果表明: 各居群的传粉者组成、主要传粉者类型及其大小各不相同, 杠杆状雄蕊及相关花部性状大小在不同居群间具有显著差异; 各居群均表现出腹部传粉和背部传粉2种传粉模式, 但背部传粉仍然是最有效的传粉方式; 居群间杠杆状雄蕊长度与传粉者体长表现出极显著的正相关, 然而花冠长与传粉者体长表现出负相关; 花冠口高度和柱头高度与传粉者胸厚也表现出一定的协同变异。鼠尾草属植物的杠杆状雄蕊及相关花部性状在传粉系统的进化过程中表现出高度的可塑性, 表明雄蕊杠杆传粉机制对传粉环境的变异非常敏感, 在该属植物的物种分化过程中具有关键作用。     

Hummingbird pollination in Salvia haenkei (Lamiaceae) lacking the typical lever mechanism

While in most Salvia species pollen is transferred by the `staminal lever mechanism', in some species the `levers' are inactive. This is also found in the bird pollinated S. haenkei from Bolivia. To understand pollen transfer in a species lacking the lever mechanism we carried out field investigations and confirmed our observations by means of morphometric measurements of both the flowers and museum skins of the observed hummingbird species. The tubular corolla forces the birds (Sappho sparganura, Colibri coruscans, Patagona gigas, Oreotrochilus adela) into a specific position thereby causing pollen transfer from the exserted pollen-sacs to the bird's feathers and bills. The staminal levers are well developed but cannot be moved because the sterile arms are closely attached to the upper face of the corolla leaving no space for any movement. We assume that the reduction of the lever mechanism reflects an adaptation to bird pollination.     

Honeyeaters and the New Zealand forest flora: The utilisation and profitability of small flowers

New Zealand flowers are frequently considered unspecialised allowing easy access to pollen and nectar by a wide range of visitors. Most conform with a syndrome of insect pollination (entomophily). Pollination of forest flowers by birds has been described for a range of species whose flowers are morphologically ornithophilous. On Kapiti Island and Little Barrier Island, all three species of New Zealand honeyeaters have been described feeding on flowers currently assumed to be entomophilous or where the pollination system is unknown. The persistence and regularity of visits suggests that the birds are obtaining suitable rewards in the form of nectar and could be serving as pollinators. We measured the nectar energetic value from flowers of three ornithophilous and five entomophilous species. Nectar production over 24 hours was highest in ornithophilous species, but the standing crop of nectar overlapped—ornithophilous species: Metrosideros fulgens (standing crop 6.6 J), Metrosideros excelsa (22 J), and Fuchsia excorticata (1.8 J); and entomophilous: Pittosporum crassifolium (23 J), Pseudopanax arboreus (1.5 J), Dysoxylum spectabile (3.7 early flowers -6.7 J late flowers), Pittosporum eugenioides (2.7 J) and Geniostoma rupestre (1.8 J). The entomophilous species present the flowers in aggregation and as result birds can visit a large number flowers per minute. We found that the average estimated nectar consumption rate for all the entomophilous species except G. rupestre was enough to sustain the two smaller New Zealand honeyeaters (hihi energy requirements= 0.12 kJ min(-1), median energy obtained: 0.16 kJ min(-1) D. spectabile—0.57 kJ min(-1) P. crassifolium); bellbird energy requirements = 0.10 kJ min(-1), median energy obtained: 0.14 kJ min(-1) D. spectabile—0.68 kJ min(-1) P. crassifolium). However, we estimate that if the birds are able to selectively forage on the flowers with most nectar, the energetic returns of all species may be sufficient for hihi and bellbird (hihi: 0.18 kJ min(-1) G. rupestre—0.93 kJ min(-1); P. crassifolium; bellbird: 0.12 kJ min(-1) G. rupestre 1.11 kJ min(-1) P. crassifolium). If tui (energy requirements: 0.25 kJ min(-1), forages randomly, only P. crassifolium (0.80 kJ min(- 1)) and D. spectabile late in the season (0.30 kJ min(-1)) provide sufficient returns, but if selective, P. arboreus (0.45 kJ min(-1)) may also suffice. We suggest that because (a) the nectar produced by entomophilous flowers provides sufficient energy to sustain the energetic requirements of birds, and (b) these plants flower in the cooler months when insect activity is reduced, birds might have played a wider role in pollination than previously considered. This finding is of particular importance because the abundance of New Zealand honeyeaters on the mainland has decreased considerably since human colonisation and this could be affecting forest regeneration.     

Functional implications of the staminal lever mechanism in Salvia cyclostegia (Lamiaceae)

Background and Aims

Flower morphology and inflorescence architecture affect pollinator foraging behaviour and thereby influence the process of pollination and the reproductive success of plants. This study explored possible ecological functions of the lever-like stamens and the floral design in Salvia cyclostegia.

Methods

Flower construction was experimentally manipulated by removing either the lower lever arms or the upper fertile thecae of the two stamens from a flower. The two types of manipulated individuals were intermixed with the control ones and randomly distributed in the population.

Key Results

Removing the sterile lower lever arms significantly reduced handling time per flower of the main pollinator, Bombus personatus. Interestingly, this manipulation did not increase the number of flowers probed per plant visit, but instead reduced it, i.e. shortened the visit sequence of the bumble-bees. Both loss of staminal lever function by removing lower lever arms and exclusion of self pollen by removing upper fertile thecae significantly reduced seed set per flower and seed set per plant. Both the manipulations interacted significantly with inflorescence size for the effect on female reproductive output.

Conclusions

Though the intact flowers demand a long handling time for pollinators, the reversible staminal lever is of advantage by promoting dispersal of pollen and thus the male function. The particular floral design in S. cyclostegia contributes to the floral constancy of B. personatus bumble-bees, with the lower lever arms acting as an optical cue for foraging cognition.     

Nectar dynamics and reproductive success in Saponaria officinalis (Caryophyllaceae) in southern Germany

Nectar production and flower visitors of the night-flowering Saponaria officinalis L. (Caryophyllaceae) were studied in relation to the reproductive success. Nectar production was worthwhile for nocturnal flower visitors. Nectar standing crop was about 267 μg sugar per flower, and comparison of nectar offering of covered and freely exposed flowers revealed that main nectar secretion time is mainly during the night up to the morning hours. In both covered and freely exposed flowers nectar volumes decreased over the day. In covered flowers, nectar volume, sugar concentration, and sugar amount per flower increased up to the third day; in older flowers sugar secretion ceased. In 1996 Autographa gamma (Noctuidae) was the exclusive nocturnal flower visitor, but pollen transfer experiments proved that A. gamma (Noctuidae) is a very ineffective pollinator of S. officinalis. In 1999 up to 50% of the observed visitors were Sphingidae, which resulted in a significantly higher seed set. Fruit set was constantly high independent of pollinator availability. In the nectar manipulation experiments seed set was highest in non-emasculated flowers filled with unnaturally high concentrated sucrose solutions. Differences to seed set on stalks treated with a sucrose solution mimicking naturally concentrated nectar were significant. Lowest fruit and seed set were found on inflorescences with emasculated flowers filled with a sucrose solution mimicking naturally concentrated nectar.     

Pollination biology in Roepera (Zygophyllaceae): How flower structure and shape influence foraging activity

The foraging behavior of bees is a complex phenomenon that depends on numerous physical features of flowers. Of particular importance are accessibility of floral rewards, floral proportions, symmetry and orientation. The flowers of Roepera are characterized by the presence of staminal scales (SS), which play an important role in nectar protection. We studied two species of Roepera with different symmetry and flower orientation, which are mainly visited by honeybees (Apis mellifera). We aimed to show how the foraging behavior of honey bees is affected by the function of SS, floral symmetry and orientation. The foraging behavior was documented by video photography. Handling time, access to nectar, percentage of pollen/nectar foraging, percentage of pollen contact and pollen deposition site on the honey bee's body were assessed. The morphometric features of the honey bees and flowers were analyzed. We found that the SS restricted pollinator access to nectar. Our results indicated consistency of visitation patterns in zygomorphic, laterally oriented flowers of R. fuscata versus random patterns in actinomorphic, diversely oriented flowers of R. leptopetala. The relative proportions of SS and proboscis length appear to be crucial for the success of pollinators. The directionality of the honey bees' movement, together with the different positioning of reproductive organs, plays an important role in the accuracy of pollen transfer and pollination efficiency.     

Correction to: The floral biology and the role of staminal connective appendages during pollination of the endoparasite Bdallophytum americanum (Cytinaceae)

Bdallophytum americanum (Cytinaceae) is an endoparasitic plant species, meaning only the flowers emerge from the host during the reproductive season. Reports on the pollination biology of this species state that its primary pollinators are carrion flies attracted by the smell of the flowers and nectar as a reward. However, the functional role of one of the most outstanding attributes of B. americanum has been neglected. These are the staminal appendages formed by the apical overgrowth of connective tissue during anther development. To determine whether these staminal appendages play a role in pollination, we monitored a nectarless population of B. americanum. We described the inflorescence emergence, floral movements, and pollination and performed field experiments to test whether the absence of the staminal connective appendages affected the visitation frequency. Male inflorescences emerge early, and both male and female flowers open during the day and do not close. Hoverflies are the most frequent visitors to both floral sexes and carry the most pollen. Moreover, the movement of staminal appendages matching the pollen viability changes is reported for the first time. The staminal appendages are the structures where pollinators land before foraging. The field experiments showed that the visitation frequency decreased sharply without staminal appendages. As a landing platform, the staminal connective appendages in B. americanum are crucial for pollinator positioning and collecting viable pollen.

     

The Staminal Lever Mechanism in Salvia L. (Lamiaceae) - a Review

Abstract: The genus Salvia encompasses about 900 species distributed world-wide. It is characterized by the famous staminal lever mechanism of the flower which is one of the best known examples of a nototribic pollination mechanism. We hypothesize that structure and functioning of the staminal levers play a major role as key structures in speciation. To cope with the complex evolutionary processes involved, a number of different methodological approaches are needed. The present paper summarizes the literature referring to structural and functional diversity, breeding systems, systematics and evolution in Salvia.     

Reproductive biology and effect of nectar robbing on fruit production in Macleania bullata (Ericaceae)

Hummingbird-pollinated flowers are frequently subjected to nectar robbing. In this paper, I examine the impact of nectar robbing on plant reproductive success on a hummingbird-pollinated species. After studying the basic aspects of the floral morphology and reproduction of Macleania bullata (Ericaceae) in a tropical montane wet forest in southwest Colombia, I examined the percent of flowers robbed and the effect of nectar robbery on fruit set. The flowers of this species are typical for plants pollinated by long-bill hummingbirds. They are protandrous and open for four days. Fruit production requires a pollinator visit; fruit set following pollinator exclusion was zero. Fruit set following xenogamous pollen transfer (36.8%) differed significantly from that of population controls (11.9%) and of autogamous pollen transfer (6.3%). Nectar volume, sugar concentration and sugar production were measured at daily intervals from bud opening until the fading of flowers. Daily nectar production (both volume and amount of sugar) varied considerably with flower age. Sugar production peaked on the second day, coinciding with the male phase. The frequency of nectar robbing in the studied population was very high (75% of examined flowers) and was positively correlated with reduced fruit set. I discuss the probability of a relation between reduced fruit set on robbed flowers and an energetic investment. Robbing by non-pollinating visitors can suppose the plant to re-synthesize more nectar. The high incidence of nectar robbing impugns the advantage of specialization.     

Staminal hairs enhance fecundity in the pollen‐rewarding self‐incompatible lily Bulbine abyssinica

Flowering plants typically use floral rewards to attract animal pollinators. Unlike nectar, pollen rewards are usually visible and may thus function as a signal that influences landing decisions by pollen‐seeking insects. Here we artificially manipulate the presence of both pollen and staminal hairs (a putative false signal of pollen reward availability) in the hermaphroditic lily Bulbine abyssinica (Xanthorrhoeaceae) to investigate their effects on bee visitation and fecundity, and also test for trade‐offs between pollen production and seed production. Honeybees, the primary floral visitors, are probably not able to distinguish between colours of petals, staminal hairs and pollen of B. abyssinica, according to analysis of reflectance spectra in a bee vision model. Flowers with both pollen and hairs removed had the lowest levels of bee visitation, seed set and seed abortions. Flowers containing hairs had an ～50% increase in visitation rate and seed set compared with emasculated flowers, while intact controls had the highest seed abortion rate. Ovule discounting in intact flowers is probably due to ovarian self‐incompatibility (or strong early inbreeding depression) as ovules penetrated by tubes from self‐pollen uniformly failed to develop into seeds. These results show that staminal hairs can enhance plant fecundity by increasing attraction of pollen‐seeking insects to flowers without increasing the risk of ovule discounting through pollinator‐mediated self‐pollination. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 481–490.     

Response of plant-pollinator communities to fire: changes in diversity, abundance and floral reward structure

Globally, plant‐pollinator communities are subject to a diverse array of perturbations and in many temperate and semi‐arid systems fire is a dominant structuring force. We present a novel and highly integrated approach, which quantifies, in parallel, the response to fire of pollinator communities, floral communities and floral reward structure. Mt Carmel, Israel is a recognised bee‐flower biodiversity hotspot, and using a chronosequence of habitats with differing post‐fire ages, we follow the changes in plant‐pollinator community organisation from immediately following a burn until full regeneration of vegetation. Initially, fire has a catastrophic effect on these communities, however, recovery is rapid with a peak in diversity of both flowers and bees in the first 2 years post‐fire, followed by a steady decline over the next 50 years. The regeneration of floral communities is closely matched by that of their principal pollinators. At the community level we quantify, per unit area of habitat, key parameters of nectar and pollen forage known to be of importance in structuring pollinator communities. Nectar volume, nectar water content, nectar concentration and the diversity of nectar foraging niches are all greatest immediately following fire with a steady decrease as regeneration proceeds. Temporal changes in energy availability for nectar, pollen, total energy (nectar+pollen) and relative importance of pollen to nectar energy show a similar general decline with site age, however, the pattern is less clear owing to the highly patchy distribution of floral resources. Changes in floral reward structure reflect the general shift from annuals (generally low‐reward open access flowers) to perennials (mostly high‐reward and restricted access flowers) as post‐fire regeneration ensues. The impact of fire on floral communities and their associated rewards have clear implications for pollinator community structure and we discuss this and the role of other disturbance factors on these systems.     

Floral bagging differentially affects handling behaviours and single-visit pollen deposition by honey bees and native bees

1. Measurements of pollinator performance are crucial to pollination studies, enabling researchers to quantify the relative value of different pollinator species to plant reproduction. One of the most widely employed measures of pollinator performance is single-visit pollen deposition, the number of conspecific pollen grains deposited to a stigma after one pollinator visit. To ensure a pollen-free stigma, experimenters must first bag flowers before exposing them to a pollinator. 2. Bagging flowers, however, may unintentionally manipulate floral characteristics to which pollinators respond. In this study, we quantified the effect of bagging on nectar volume in watermelon (Citrullus lanatus) flowers, and how this affects pollinator performance and behaviour. 3. Experimental bagging resulted in roughly 30-fold increases in nectar volume relative to unmanipulated, open-pollinated field flowers after only a few hours. Honey bees, but not native bees, consistently displayed elevated handling times and single-visit pollen deposition on unmanipulated bagged flowers relative to those from which we removed nectar to mimic volumes in open-pollinated flowers. 4. Furthermore, we identify specific bee foraging behaviours during a floral visit that account for differences in pollen deposition, and how these differ between honey bees and native bees. 5. Our findings suggest that experimental bagging of flowers, without accounting for artificially accumulated nectar, can lead to biased estimates of pollinator performance in pollinator taxa that respond strongly to nectar volume. We advise that pollination studies be attentive to nectar secretion dynamics in their focal plant species to ensure unbiased estimates of pollinator performance across multiple pollinator species.     

Nectar Yeasts in the Tall Larkspur Delphinium barbeyi (Ranunculaceae) and Effects on Components of Pollinator Foraging Behavior

Microorganisms frequently colonize the nectar of angiosperm species. Though capable of altering a suite of traits important for pollinator attraction, few studies exist that test the degree to which they mediate pollinator foraging behavior. The objective of our study was to fill this gap by assessing the abundance and diversity of yeasts associated with the perennial larkspur Delphinium barbeyi (Ranunculaceae) and testing whether their presence affected components of pollinator foraging behavior. Yeasts frequently colonized D. barbeyi nectar, populating 54–77% of flowers examined depending on site. Though common, the yeast community was species-poor, represented by a single species, Metschnikowia reukaufii. Female-phase flowers of D. barbeyi were more likely to have higher densities of yeasts in comparison to male-phase flowers. Pollinators were likely vectors of yeasts, as virgin (unvisited) flowers rarely contained yeasts compared to flowers open to pollinator visitation, which were frequently colonized. Finally, pollinators responded positively to the presence of yeasts. Bombus foragers both visited and probed more flowers inoculated with yeasts in comparison to uninoculated controls. Taken together, our results suggest that variation in the occurrence and density of nectar-inhabiting yeasts have the potential to alter components of pollinator foraging behavior linked to pollen transfer and plant fitness.     

Bird pollination syndrome is the plant's adaptation to ornithophily,but nectarivorous birds are not so selective

Many tropical plants are pollinated by birds and several bird phylogenetical lineages have specialised to a nectar diet. The long-assumed, intimate ecological and evolutionary relationship between ornithophilous plants and phenotypically specialised nectarivorous birds has nevertheless been questioned in recent decades, where such plant–pollinator interactions have been shown to be highly generalised. In our study, we analysed two extensive interaction datasets: bird–flower and insect–flower interactions, both collected on Mt Cameroon, west-central Africa. We tested if: 1) insects and birds interact with distinct groups of plants; 2) plants with a typical set of ornithophilous floral traits (i.e. bird pollination syndrome) interact mainly with birds; 3) birds favour plants with bird pollination syndrome and; 4) if and how the individual floral traits and plant level nectar production predict bird visitation. Bird-visited plants were typically also visited by insects, while approximately half of the plants were visited by insects only. We confirmed the validity of the bird pollination syndrome hypothesis, as plants with bird-pollination syndrome traits were visited by birds at a higher rate and mostly hosted a lower frequency of visiting insects. However, these ornithophilous plants were not more attractive than the other plants for nectar-feeding birds. Nectar production per plant individual was a better predictor of bird visitation than any other floral trait traditionally related to the bird pollination syndrome. Our study thus demonstrated the highly asymmetrical relationship between ornithophilous plants and nectarivorous birds.     

The effects of nectar addition on pollen removal and geitonogamy in the non-rewarding orchid Anacamptis morio

It has been suggested that the absence of floral rewards in many orchid species causes pollinators to probe fewer flowers on a plant, and thus reduces geitonogamy, i.e. self-pollination between flowers, which may result in inbreeding depression and reduced pollen export. We examined the effects of nectar addition on pollinator visitation and pollen transfer by tracking the fate of colour-labelled pollen in Anacamptis morio, a non-rewarding orchid species pollinated primarily by queen bumble-bees. Addition of nectar to spurs of A. morio significantly increased the number of flowers probed by bumble-bees, the time spent on an inflorescence, pollinarium removal and the proportion of removed pollen involved in self-pollination through geitonogamy, but did not affect pollen carryover (the fraction of a pollinarium carried over from one flower to the next). Only visits that exceeded 18 s resulted in geitonogamy, as this is the time taken for removed pollinaria to bend into a position to strike the stigma. A mutation for nectar production in A. morio would result in an initial 3.8-fold increase in pollinarium removal per visit, but also increase geitonogamous self-pollination from less than 10% of pollen depositions to ca. 40%. Greater efficiency of pollen export will favour deceptive plants when pollinators are relatively common and most pollinaria are removed from flowers or when inbreeding depression is severe. These findings provide empirical support both for Darwin's contention that pollinarium bending is an anti-selfing mechanism in orchids and for the idea that floral deception serves to maximize the efficiency of pollen export.     

Staminal evolution in the genus Salvia (Lamiaceae): molecular phylogenetic evidence for multiple origins of the staminal lever

BACKGROUND AND AIMS: The genus Salvia has traditionally included any member of the tribe Mentheae (Lamiaceae) with only two stamens and with each stamen expressing an elongate connective. The recent demonstration of the non-monophyly of the genus presents interesting implications for staminal evolution in the tribe Mentheae. In the context of a molecular phylogeny, the staminal morphology of the various lineages of Salvia and related genera is characterized and an evolutionary interpretation of staminal variation within the tribe Mentheae is presented. METHODS: Two molecular analyses are presented in order to investigate phylogenetic relationships in the tribe Mentheae and the genus Salvia. The first presents a tribal survey of the Mentheae and the second concentrates on Salvia and related genera. Schematic sketches are presented for the staminal morphology of each major lineage of Salvia and related genera. KEY RESULTS: These analyses suggest an independent origin of the staminal elongate connective on at least three different occasions within the tribe Mentheae, each time with a distinct morphology. Each independent origin of the lever mechanism shows a similar progression of staminal change from slight elongation of the connective tissue separating two fertile thecae to abortion of the posterior thecae and fusion of adjacent posterior thecae. A monophyletic lineage within the Mentheae is characterized consisting of the genera Lepechinia, Melissa, Salvia, Dorystaechas, Meriandra, Zhumeria, Perovskia and Rosmarinus. CONCLUSIONS: Based on these results the following are characterized: (1) the independent origin of the staminal lever mechanism on at least three different occasions in Salvia, (2) that Salvia is clearly polyphyletic, with five other genera intercalated within it, and (3) staminal evolution has proceeded in different ways in each of the three lineages of Salvia but has resulted in remarkably similar staminal morphologies.     

The assembly of plants used as nectar sources by hummingbirds in a Cerrado area of Central Brazil

Studies on hummingbird–plant interactions commonly use a pollination approach emphasizing mutualistic relationships. But floral resources are often used opportunistically by these birds. Plant–pollinator assemblies and pollination sustainability will depend both on the well-adapted plants and other potential floral resources. The Cerrado, Neotropical savannas of Central Brazil, has ca. 7.5 % of its flora supposedly adapted to hummingbird pollination. But detailed information about flowers effectively used by hummingbirds at community level is still lacking. Hence, we recorded all plant species visited by hummingbirds, to determine how these nectariferous flowers were distributed in time and space in different plant formations of a Cerrado area, and also the hummingbird species that visit them. The study was conducted between March 2007 and December 2008 in the Panga Ecological Station. Data regarding flowering phenology, floral morphology and visitation were collected monthly. Forty-six nectariferous species from 39 genera and 17 families were recorded, most with annual flowering dynamics and tubular flowers. But only 21 species had a combination of traits fitting classic ornithophilous syndrome. For the remaining species hummingbird visitation was ascertained from observations at the study site or other sites in the region. Eight hummingbird species occurred in the area and were recorded visiting directly 36 plant species. The study area presented a relatively low number of ornithophilous plants, but a great habitat diversity and many non-ornithophilous plants that hummingbirds used as nectar sources. Therefore, in the studied Cerrado, the diversity of environments and nectariferous plants favour the maintenance of resident and migrant hummingbirds.