Hummingbird responses to gender-biased nectar production: are nectar biases maintained by natural or sexual selection?

Pollinators mediate the evolution of secondary floral traits through both natural and sexual selection. Gender-biased nectar, for example, could be maintained by one or both, depending on the interactions between plants and pollinators. Here, I investigate pollinator responses to gender-biased nectar using the dichogamous herb Chrysothemis friedrichsthaliana (Gesneriaceae) which produces more nectar during the male floral phase. Previous research showed that the hummingbird pollinator Phaethornis striigularis visited male-phase flowers more often than female-phase flowers, and multiple visits benefited male more than female fecundity. If sexual selection maintains male-biased rewards, hummingbirds should prefer more-rewarding flowers independent of floral gender. If, however, differential rewards are partially maintained through natural selection, hummingbirds should respond to asymmetry with visits that reduce geitonogamy, i.e. selfing and pollen discounting. In plants with male biases, these visit types include single-flower visits and movements from low to high rewards. To test these predictions, I manipulated nectar asymmetry between pairs of real or artificial flowers on plants and recorded foraging behaviour. I also assessed maternal costs of selfing using hand pollinations. For plants with real flowers, hummingbirds preferred more-rewarding flowers and male-phase morphology, the latter possibly owing to previous experience. At artificial arrays, hummingbirds responded to extreme reward asymmetry with increased single-flower visits; however, they moved from high to low rewards more often than low to high. Finally, selfed flowers did not produce inferior seeds. In summary, sexual selection, more so than geitonogamy avoidance, maintains nectar biases in C. friedrichsthaliana, in one of the clearest examples of sexual selection in plants, to date.     

Flower morphology,nectar traits and pollinators of Cerinthe major (Boraginaceae-Lithospermeae)

In April and May 2010, a natural population of Cerinthe major (Boraginaceae-Lithospermeae) was investigated with regard to: floral morphology, phenology, sexual receptivity during anthesis, the production of nectar and its sugar and amino acid content. The pollinator array of this species was also investigated. Cerinthe major has showy, pentamerous, hermaphroditic flowers with a deep yellow corolla tube arranged on numerous scorpioid inflorescences. Nectar and pollen are the main floral rewards for pollinators. The arrangement of stamens limits access to nectar and promotes pollen loading onto pollinator bodies as they force their way towards the source of nectar. By limiting the direct exposure of nectar to the environment, the corolla tube and the arrangement of anther filaments seemingly protect nectar from evaporation resulting from high temperatures and low relative humidity during flower anthesis. The green, annular nectary located at the base of the ovary lobes is composed of a very thin epidermis enclosing a dense mass of parenchyma. The epidermis lacks stomata, and the thin cuticle has an irregular surface. The parenchyma cells do not store starch during the pre-secretory stage, suggesting that immediate photosynthesis is the most likely source of nectar carbohydrates. Generally, anthesis lasts 4–5 days; the gynoecium becomes fully receptive on the second day and this is synchronous with anther dehiscence. Since nectar production begins during the bud stage, a substantial volume is available for flower visitors at anthesis. Nectar production decreases drastically in senescent flowers and, if not consumed, the nectar can be re-absorbed. Analysis of sugars revealed a sucrose-dominant nectar (sucrose = 93.18 ± 1.35%). Proline, GABA, taurine, leucine, citrulline and alanine were the main amino acids present and are probably implicated in pollinator flight performance.     

草乌花蜜产量的梯度分布及熊蜂自下而上的访花行为

收益降低假说(declining reward hypothesis)认为熊蜂自下而上的访花顺序是对花蜜产量的直接响应,先访问下部花蜜产量高的花可以获得更多的收益;花开口方向假说认为自下而上访花是因为熊蜂更容易看见其上部的花朵。为了验证上述两个假说,我们于2008年8月在北京小龙门国家森林公园调查了红光熊蜂(Bombus ignitus)访问草乌(Aconitum kusnezoffii)直立和倒立顶生花序的访问顺序,测量了直立花序下部雌性阶段花和上部雄性阶段花花蜜的糖浓度、体积,计算了花蜜中的糖含量。结果表明,红光熊蜂在直立花序和倒立花序内均以向上运动为主,分别占总运动次数的62.77%和68.35%;直立花序下部雌性阶段花花蜜糖浓度比上部雄性阶段花低1.44%,但是花蜜体积和花蜜中的糖含量都显著高于雄性阶段的花。由于熊蜂访问倒立花序时先访问的是下部的低回报雄性阶段的花,然后再访问上部高回报的雌性阶段的花,这与收益降低假说矛盾,表明红光熊蜂自下而上访问草乌直立花序可能不是受到花蜜产量的调节。     

Selection for floral integration and trait variation in zygomorphic flowers of Aconitum japonicum ssp. subcuneatum (Ranunculaceae)

Pollinator-mediated selection might lead to among-trait differences in the degree and pattern of floral integration and intra-flower variation. To examine the patterns of intra-flower variation in floral traits, including nectar volume, we performed a field study using the zygomorphic flowers of Aconitum japonicum ssp. subcuneatum. We investigated (1) correlations between the sizes of the left and right sepals and petals, (2) variation in floral traits among plants, within plants and within flowers, (3) effects of sexual phases on floral integration variation in floral and nectar traits, and (4) the effect of size and intra-flower variation in traits of the left and right sepals and petals on pollen removal by pollinators. Lateral sepal area, but not lower sepal area, was highly correlated between the left and right sepals. Floral traits were more integrated during the male phase than during the female phase. Nectar standing crop in male-phase flowers correlated with helmet height and lateral and lower sepal area, but in female-phase flowers it only correlated with spur length. While intra-flower variance in lateral sepal area accounted for approximately 10% of the overall variance in these traits, the variance in lower sepal area accounted for 70% of the overall variance. Lateral sepal area had a negative effect on the number of pollen grains remaining after pollinator visits. Low variance in lateral sepals within flowers and measurements of pollen removal suggest that lateral sepals play a more important role in pollen export than the other traits. Left and right sepals may be the targets of selection for symmetry in zygomorphic flowers.

     

Nectar ecology of Datura ferox (Solanaceae): an invasive weed with nocturnal flowers in agro-ecosystems from central Argentina

Plant–pollinator interactions provide highly important ecological functions, and are influenced by floral nectar characteristics. The night blooming Datura ferox is an excellent model to test general hypotheses on the relationship between nectar traits (e.g., nectar secretion patterns, nectar chemical composition), pollinators and reproductive success for invasive, weedy species in highly modified ecosystems as crop fields. We hypothesized an adjustment between nectar composition and secretion dynamics through flower anthesis and the activity and requirements of nocturnal pollinators. Nectar chemical analyses showed low quantities of amino acids and lipids, phenolics, and alkaloids were not detected. D. ferox showed sucrose-dominant nectar with comparable amount of hexoses. Sugar proportions did not vary between populations or during flowering season. Most nectar is secreted before flower opening. Nectar resorption was detected at the end of anthesis. Experimentally drained flowers of both populations increased nectar production up to 50 % in the total amount of sugar per flower compared to control flowers. Nectar standing crop was relatively constant during the flowering season, but differences were detected between populations. Nectar traits of D. ferox would be favoring cross-pollination and maintaining seed production of this weed, since recently open flowers display a higher amount of nectar and they can renew nectar after a pollinator visit or reabsorb it at the end of anthesis. This nectar source may be important for native pollinators considering that human-induced forest fragmentation is related with the impoverishment of native flora from agro-ecosystems.     

Male-biased nectar production in a protandrous herb matches predictions of sexual selection theory in plants

Nectar production may disproportionately benefit male relative to female pollination success. In such cases, sexual selection is often suggested as the cause of asymmetric benefits, yet sexual selection in plants-particularly plants with hermaphroditic flowers-is infrequently tested empirically. Here, I used a protandrous herb with male-biased nectar production (Chrysothemis friedrichsthaliana, Gesneriaceae) to test predictions from sexual selection theory. During three flowering seasons, I measured nectar production, pollinator visits, and male and female fecundity following different numbers of cross-pollination events. In accordance with sexual selection predictions, (1) nectar production was greater during the male phase by at least 65%; (2) visits by the main pollinator (hummingbird Phaethornis striigularis) were limiting for part of the season, indicating that plants had to compete for pollinator visits; (3) pollinators spent 53% more time per visit and made 86% more visits to male- vs. female-phase flowers, suggesting that nectar increased male more than female pollination success; and (4) female fecundity was maximized by one visit, whereas male fecundity continued to increase with additional visits. Autonomous self-pollination further reduced visit requirements for maximum female seed set. These findings match specific sexual selection predictions: they link an observable male bias in a secondary sexual trait (nectar) to positive responses of mating participants (pollinators), resulting in more mating opportunities for mate-limited males, relative to apparently resource-limited females. This field-testing of theoretical predictions provides unique evidence that sexual selection helps maintain nectar production patterns in this and, quite likely, other hermaphroditic plant species.     

Main sugar composition of floral nectar in three species groups of Scrophularia (Scrophulariaceae) with different principal pollinators

In some angiosperm groups, a parallelism between nectar traits and pollination syndromes has been demonstrated, whereas in others there is not such relationship and it has been explained as due to phylogenetic constraints. However, nectar trait information remains scarce for many plant groups. This paper focuses on three groups of Scrophularia species, with different flower sizes and principal pollinators, to find out whether nectar sugar composition is determined by pollinator type or reflects taxonomic affinities. Since the species we examined have protogynous flowers, and gender bias in nectar sugar composition has been noted in few plant groups, we also investigated whether sexual phase influenced Scrophularia nectar composition. The sugar composition was found to be similar in all species, having high‐sucrose nectar, except for the Macaronesian Scrophularia calliantha, which was the only species with balanced nectar; this last kind of nectar could be associated with the high interaction rates observed between S. calliantha and passerine birds. The nectar sugar composition (high in sucrose) was unrelated to the principal pollinator group, and could instead be considered a conservative taxonomic trait. No gender bias was observed between functionally female and male flowers for nectar volume or concentration. However, sexual phase significantly affected sucrose percentage in the largest‐flowered species, where the female phase flowers had higher sucrose percentages than the male phase flowers.     

Effects of variation in inflorescence size and floral rewards on the visitation rates of traplining pollinators ofAralia hispida

Summary In field experiments withAralia hispida inflorescences, the following variables were manipulated: number of umbels per inflorescence, number of flowers per umbel, and amounts of pollen and nectar per flower. Visitation rates by bumble bees, the principal pollinators, were then observed. In the reward-variation experiments, bees appeared to learn the positions of nectar-rich shoots, and visited them significantly more often than nectar-poor shoots. They did not respond to similar variation in pollen production. The nectar preferences developed slowly after the treatments were imposed, and bees continued to favor sites that had been occupied by nectar-rich shoots even after the treatments were discontinued. Visitation rate was approximately proportional to flower number, making it unlikely that increases in inflorescence size produced a disproportionate gain in male reproductive success (a necessary condition in certain models for the evolution of dioecy). For a fixed number of flowers per inflorescence, bees preferred inflorescences with more umbels. In pairwise choice tests of male-phase and female-phase umbels of various sizes, bees preferred male-phase umbels and larger umbels; the preference for male-phase umbels is stronger in bees that had previously fed on male-phase umbels.     

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.     

Differences in the diversity and composition of the pollinator assemblage of two co-flowering congeneric alpine wallflowers, Erysimum nevadense and E. baeticum

Many alpine plants display a generalist pollination system where almost any available flower visitor could act as a legitimate pollinator. Co-flowering related plant species displaying a generalized pollination system can decrease their fitness due to interspecific mating. In this study, we determine the difference in diversity and composition of the pollination systems of two co-occurring species endemic to the alpine region of the Sierra Nevada (SE Spain), Erysimum nevadense Reut. and Erysimum baeticum baeticum (Heywood) Polatschek (Brassicaceae), and check for the potential role of floral and plant traits in explaining the observed differences. For this, we labeled 30 plants in two populations of each plant species located in the same area. We determined flower visitor assemblage by sampling all insects approaching the flowers and contacting the sexual organs during 60-min surveys. We found that the diversity of the pollinator assemblage was similar between the two studied wallflower species, both species showing a much generalized pollination system. However, the composition of the pollinator assemblage was different, since the flowers of E. nevadense were visited mostly by beetles and in a lesser extent by hover-flies, whereas the flowers of E. baeticum baeticum were visited mostly by ants and bees. Interestingly, flower traits varied between species, with E. nevadense displaying yellow, smaller and shallower flowers and E. baeticum baeticum displaying purple, large and deeper flowers. These findings suggest that differences in floral traits can explain the observed differences in the composition of the pollinator assemblage between both wallflower species.     

Pollen and nectar as a reward in the basal epidendroid Psilochilus modestus (Orchidaceae: Triphoreae): A study of floral morphology, reproductive biology and pollination strategy

Psilochilus modestus Barb. Rodr. is a basal epidendroid orchid occurring in both the semi-deciduous and Atlantic rain forests of the state of São Paulo, southeastern Brazil. This species presents a perfect flowering synchrony within populations, since all the mature buds of each plant open simultaneously in the morning hours of the same day. These flowers are available only for 1 day and are pollinated by several species of small solitary and social native bees. These bees exploit both pollen and nectar as a reward. The bees collecting pollen promote a higher fruit set and perform mainly self-pollination while those collecting nectar, which are less numerous, are responsible for an increase in cross-pollinations. P. modestus is self-compatible but pollinator dependent. Natural fruit set (open pollination) is low when compared with the numbers obtained under manual self- and cross-pollination. Low fruit set in natural conditions is related to deficient pollen transfer, and pollinator inefficiency seems to be the main factor. Some factors, such as the small amount of nectar produced, the low number of flowers per inflorescence and their availability for 1 day only added to the perfect flowering synchrony seem to be responsible for the increase of cross-pollinations. The offering of both pollen and nectar as a reward can represent a transitional condition in basal Epidendroideae. Based on floral morphology, reward production and pollinator behavior, the relationship of P. modestus with the basal and most derived groups within Orchidaceae is discussed.     

Spatial and temporal patterns of floral scent emission in Dianthus inoxianus and electroantennographic responses of its hawkmoth pollinator

Scent emission is important in nocturnal pollination systems, and plant species pollinated by nocturnal insects often present characteristic odor compositions and temporal patterns of emission. We investigated the temporal (day/night; flower lifetime) and spatial (different flower parts, nectar) pattern of flower scent emission in nocturnally pollinated Dianthusinoxianus, and determined which compounds elicit physiological responses on the antennae of the sphingid pollinator Hyles livornica.The scent of D.inoxianus comprises 68 volatile compounds, but is dominated by aliphatic 2-ketones and sesquiterpenoids, which altogether make up 82% of collected volatiles. Several major and minor compounds elicit electrophysiological responses in the antennae of H. livornica. Total odor emission does not vary along day and night hours, and neither does along the life of the flower. However, the proportion of compounds eliciting physiological responses varies between day and night. All flower parts as well as nectar release volatiles. The scent of isolated flower parts is dominated by fatty acid derivatives, whereas nectar is dominated by benzenoids. Dissection (= damage) of flowers induced a ca. 20-fold increase in the rate of emission of EAD-active volatiles, especially aliphatic 2-ketones.We suggest that aliphatic 2-ketones might contribute to pollinator attraction in D. inoxianus, even though they have been attributed an insect repellent function in other plant species. We also hypothesize that the benzenoids in nectar may act as an honest signal (‘nectar guide’) for pollinators.     

The reproductive biology ofJaborosa integrifolia (Solanaceae): Why its fruits are so rare?

Jaborosa integrifolia exhibits stigma-height polymorphism. There are individuals with flowers where anthers and stigma are at the same height but the rule is variable herkogamy, the most common type (75%) being that with an exerted stigma. Self- and cross-tubes did not differ in their capability to reach the ovary (t = –0.67,P < 0.53); they had a high growth rate (6.95 ± 2.28 mm h^–1). There is not autogamy but mostly self-incompatibility. Fruits from controlled cross-pollination showed the highest seed set and seed viability. The nectar sugar is characterized by a similar amount of glucose and fructose, and by the absence of sucrose. Although nectar secretion was continuous throughout the life of the flower, most nectar was secreted during the first 24 h after flower opening. Nectar production costs appear to be lower than in other species since nectar secretion is neither inhibited after a removal (i.e. a pollinator visit) nor reabsorbed as the flower ages. Sphingids visit the flowers mainly after midnight. They insert their proboscis down to the base of the corolla tube to reach the nectar. The upper limit to fruit production is set by pollinator visits. Fruits produced from open-pollinated flowers are often predated by numerous larvae (mainly lepidopteran ones). Considering that this species is mostly self-incompatible and pollination is limited, that each plant displays only a low number of flowers throughout the flowering season, and that there is a high rate of fruit predation, it is not surprising that fruits ofJ. integrifolia are so rare.     

FLORAL NECTAR SUGAR COMPOSITION AND POLLINATOR TYPE AMONG NEW WORLD GENERA IN TRIBE ANTIRRHINEAE (SCROPHULARIACEAE)

The floral nectar sugar compositions of 20 New World species from 10 genera and of five interspecific hybrids in tribe Antirrhineae have been analyzed using high-performance liquid chromatography. Species are pollinated by short-tongued bees, long-tongued bees, and hummingbirds. Ornithophily represents the derived condition in the tribe and has arisen independently in subtribes Maurandyinae and Gambeliinae. All nectars analyzed are sucrose-dominant or -rich, except for the hexose-rich nectar of Mohavea breviflora. Despite the predominance of sucrose, floral nectar sugars from species pollinated by different pollen vectors have characteristic constituents. Nectar sugars from flowers visited by hummingbirds average 76.2% sucrose and have compositions remarkably similar to hummingbird nectars analyzed in previous studies of unrelated species. Long-tongued bee nectars average 87% sucrose and differ from shorttongued bee nectars which have the lowest mean sucrose percentage (40.2%). The association of sugar constituent types and principal pollinators is concordant with previous data and supports hypotheses concerning pollinator preferences and the adaptive significance of certain nectar sugar compositions. Within this adaptive framework, phylogenetic constraint is also operative and may explain the predominance of sucrose in nectar sugars, similarities in sugar composition among hummingbird nectars in subtribes Maurandyinae and Gambeliinae, and the similarity of nectar from Galvezia leucantha (long-tongued bee-pollinated) to hummingbird-pollinated species also in subtribe Gambeliinae.     

Efficient avian pollination of Strelitzia reginae outside of South Africa

In its native South Africa, endemic birds pollinate the complex flowers of Strelitzia reginae (bird of paradise) through a highly complex method of pollination. The plant is cultivated worldwide in warm-temperated regions but systematic pollination of the ornithophilous species by local birds has not been reported, and, consequently, seed production is rare outside of South Africa. We found that a member of the New World warblers, Geothlypis trichas, efficiently carried out pollination of S. reginae in southern California, thereby supplementing its typical diet of insects with the energy-rich nectar of S. reginae. Only occasionally, seeds were found in plantings not visited by these birds. The pollinator service provided by the warbler increases seed production in an area outside of South Africa. This could lead to adaptive changes in the exotic species, advance species establishment and persistence and possibly promote invasive behavior in a non-native environment.     

青藏高原露蕊乌头不同海拔居群的开花持续期与繁殖分配

对青藏高原东部3个不同海拔居群的露蕊乌头(Aconitum gymnandrum)(毛茛科)开花持续期和繁殖分配进行了野外观察和检测试验。结果表明:(1)随着海拔的升高,露蕊乌头的开花持续期时间延长,且不同居群间存在显著差异。(2)露蕊乌头的植株个体越大,繁殖投入越高。(3)露蕊乌头的繁殖分配在地上总生物量中所占的比例随海拔升高而增加。研究认为,露蕊乌头在高海拔地区较高的繁殖分配比例强调了有性繁殖在高山恶劣环境中的重要性,而高海拔地区开花持续期的延长补偿了传粉昆虫的减少并增加了对传粉昆虫的吸引能力。     

Sex differential nectar secretion in protandrous Alstroemeria aurea (Alstroemeriaceae): is production altered by pollen removal and receipt?

We examined diurnal and nocturnal nectar secretion across sexual stages in protandrous Alstroemeria aurea, a bumble bee-pollinated herb with long-lived flowers native to the southern Andes. We found the following patterns: (1) most nectar was produced diurnally and (2) three times more sugar was secreted during the male than female phase, not only because the male phase lasted longer but also because the rate of nectar production was higher. This 3:1 ratio in nectar production matched the ratio of the minimum number of bumble bee visits required on average to saturate male (pollen removal) vs. female (seed set) functions. Standing crop of nectar, on the other hand, did not differ greatly between male- and female- stage flowers left open to visitors, because the high-production male-phase flowers were visited more frequently than female- phase flowers. In an experiment concurrent with the repeated nectar sampling of individual flowers over their life-span, we removed pollen from anthers or deposited pollen on stigmas by hand. Neither treatment, designed to mimic effects of visits by Alstroemeria's native bumble bee pollinator, affected nectar production. The absence of plasticity in nectar secretion in relation to pollination events may reflect a low cost of nectar production, or may result from developmental constraints related to the evolution of the synchronous protandry that characterizes A. aurea.     

Molecular and morphological analyses of EuropeanAconitum species (Ranunculaceae)

The systematics of the yellow-floweredAconitum lycoctonum species complex (Ranunculaceae) has long been considered difficult because of high morphological variability and hypothesized hybridization. To elucidate the systematic value of these taxa, and to study their phylogeny in a broader context, we studied all species of the subgenusLycoctonum from Europe and the Caucasus Mountains and the European species of the subgen.Aconitum. We sequenced the ITS region and the chloroplast intergenic spacerpsbA-trnH and conducted parsimony analyses. We also measured morphological characters from 19A. lycoctonum populations. The two subgenera in Europe are clearly separated but hybridization between subgenera might have taken place in the past. Sequence variation withinA. lycoctonum and among all taxa of subgenusLycoctonum was very low and probably the result of recent speciation, leading to one morphologically highly variable species in which color morphs could be seen as subspecies. There was high morphological variability within and among populations and the morphological characters have no value as systematic characters.     

Nectar sugar composition of European Caryophylloideae (Caryophyllaceae) in relation to flower length,pollination biology and phylogeny

Floral nectar composition has been explained as an adaptation to factors that are either directly or indirectly related to pollinator attraction. However, it is often unclear whether the sugar composition is a direct adaptation to pollinator preferences. Firstly, the lower osmolality of sucrose solutions means that they evaporate more rapidly than hexose solutions, which might be one reason why sucrose‐rich nectar is typically found in flowers with long tubes (adapted to long‐tongued pollinators), where it is better protected from evaporation than in open or short‐tubed flowers. Secondly, it can be assumed that temperature‐dependent evaporation is generally lower during the night than during the day so that selection pressure to secrete nectar with high osmolality (i.e. hexose‐rich solutions) is relaxed for night‐active flowers pollinated at night. Thirdly, the breeding system may affect selection pressure on nectar traits; that is, for pollinator‐independent, self‐pollinated plants, a lower selective pressure on nectar traits can be assumed, leading to a higher variability of nectar sugar composition independent of pollinator preferences, nectar accessibility and nectar protection. To analyse the relations between flower tube length, day vs. night pollination and self‐pollination, the nectar sugar composition was investigated in 78 European Caryophylloideae (Caryophyllaceae) with different pollination modes (diurnal, nocturnal, self‐pollination) using high‐performance liquid chromatography (HPLC). All Caryophylleae species (Dianthus and relatives) were found to have nectar with more than 50% sucrose, whereas the sugar composition of Sileneae species (Silene and relatives) ranged from 0% to 98.2%. In the genus Silene, a clear dichotomous distribution of sucrose‐ and hexose‐dominant nectars is evident. We found a positive correlation between the flower tube length and sucrose content in Caryophylloideae, particularly in day‐flowering species, using both conventional analyses and phylogenetically independent contrasts.     

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.