Phenotypic selection on floral traits in the arctic plant Parrya nudicaulis (Brassicaceae)

The evolution of floral traits is often attributed to pollinator‐mediated selection; however, the importance of pollinators as selective agents in arctic environments is poorly resolved. In arctic and subarctic regions that are thought to be pollen limited, selection is expected to either favor floral traits that increase pollinator attraction or promote reproductive assurance through selfing. We quantified phenotypic selection on floral traits in two arctic and two subarctic populations of the self‐compatible, but largely pollinator‐dependent, Parrya nudicaulis. Additionally, we measured selection in plants in both open pollination and pollen augmentation treatments to estimate selection imposed by pollinators in one population. Seed production was found to be limited by pollen availability and strong directional selection on flower number was observed. We did not detect consistently greater magnitudes of selection on floral traits in the arctic relative to the subarctic populations. Directional selection for more pigmented flowers in one arctic population was observed, however. In some populations, selection on flower color was found to interact with other traits. We did not detect consistently stronger selection gradients across all traits for plants exposed to pollinator selection relative to those in the pollen augmentation treatment; however, directional selection tended to be higher for some floral traits in open‐pollinated plants.     

Selective nectar robbing in a gynodioecious plant (Glechoma longituba) enhances female advantage

Nectar robbing not only affects the reproductive fitness of the plant but it may also potentially affect the pollination dynamics of the associated coflowering individuals. In this study, we established that the nectar robber Xylocopa sinensis robs nectar only from the hermaphrodite ramets of the gynodioecious plant Glechoma longituba but not from the female ramets. In populations with high rates of nectar robbing, this results in hermaphrodite ramets having reduced seed set whereas the female ramets have a slightly increased seed set. We hypothesize that selective nectar robbing confers an advantage to female individuals and thus ensures their maintenance in gynodioecious populations. Results of controlled experiments indicated that the reduction in the amounts of nectar available occasioned by nectar robbing resulted in some legitimate pollinators switching to visiting flowers on female rather than hermaphrodite ramets. This resulted in lower pollination rates and seed set for hermaphrodites and higher pollination rates and seed set for females. This study presents a previously unreported mechanism causing female advantage in gynodioecious plants.     

Character displacement in style length between pollinator-sharing Clerodendrum trichotomum and C. izuinsulare (Verbenaceae)

Competition for pollination may occur between pollinator-sharing sympatric plants and this may cause character displacement of their floral traits. We examined this possibility by comparing flower morphology of the sympatric population of Clerodendrum trichotomum and its co-flowering congener, C. izuinsulare, with that of the allopatric populations. The two species were visited in common by such insects as diurnal hawkmoths, bees, swallowtails and nocturnal hawkmoths, and were pollinated nocturnally as well as diurnally. Interspecific pollen transfer can occur by sharing pollinators; however, they did not hybridize when artificially pollinated. Flower size, including stamen and style lengths, is larger in C. trichotomum with an overlap in range. The style of C. izuinsulare in the sympatric population was significantly shorter than that in the allopatric population, while there was no significant difference in style length between the allopatric and the sympatric C. trichotomum. This seems to facilitate avoidance of interspecific pollen transfer in the sympatric population.Ken Inoue was killed in an accident on July 28, 2003, during his research trip in Sakhalin.     

Floral traits variation, legitimate pollination, and nectar robbing in Polygala vayredae (Polygalaceae)

The variation of floral morphology and its effect on the flower visitors of Polygala vayredae Costa (Polygalaceae), a narrow endemic species from the Oriental pre-Pyrenees, were examined. First, to account for the main floral reward (i.e., nectar), the relationship between the dimensions of the nectar gland and nectar production was investigated. Second, floral traits variation was assessed within and between the three most representative populations of the species. Finally, the role of several floral traits in the female fitness was evaluated. Furthermore, as nectar robbing was highly frequent, preferences of robbers for specific floral traits and their impact on legitimate pollinations were also evaluated. The flowers of this species are characterized by significant variations in floral characteristics and nectar rewards. A significant and positive correlation between the nectar gland dimensions and nectar production per flower was observed, with the gland dimensions being a good measure to infer the rewards offered by the flowers of P. vayredae. In general, corolla traits were significant and positively correlated with each other. Nectar was revealed to be an important trait in flower–visitor interactions, with legitimate pollinations being primarily influenced by this floral reward. Negative correlations between robbing frequency and legitimate pollinations were observed in two of the studied populations, and positive correlations between flower size and robbing frequency were observed in one population. An indirect negative selection over phenotypic floral traits mediated by nectar robbers is proposed.     

Patterns and implications of floral nectar secretion, chemical composition, removal effects and standing crop in Mandevilla pentlandiana (Apocynaceae)

Flower morphology, nectar features (chemical composition, production pattern, removal effects, and standing crop) and floral visitors are analysed in an Argentine population of Mandevilla pentlandiana. Nectar variability was examined during the lifetime of a single flower, over the course of the flowering season, and at different times of the day. Nectar is sucrose dominant. There were some variations in the proportions of sugar throughout both the flower lifespan and the flowering season. Flowers produced most nectar during bud-stage. Nectar secretion ceased near the end of the first day after flower opening. Nectar quantity varied as a function of flower age due to a combination of nectar secretion, cessation, and resorption periods. Overall sugar production was increased by nectar removal. Standing crop data showed that each open flower and inflorescence offers c. 2 and 11 mg of sugar respectively at any time of the flowering season. There was higher nectar availability at the beginning of the flowering season compared with the rest of the period. Flowers were visited by bumblebees, honeybees and hummingbirds. The greater the number of open flowers and the nectar variance, the more the mean reward quantity per flower available in the inflorescence. The sources of nectar variability in M. pentlandiana seem to be linked with both the female function (nectar resorption, nectar cessation) and the male one (early and comparatively large nectar availability, variation in nectar production as the flower ages, nectar secretion stimulation by nectar removals).     

Pollination Biology of the Exotic Rattleweed Crotalaria retusa L. (Fabaceae) in NE Brazil1

The rattleweed Crotalaria retusa was introduced in Brazil from Africa, and combines a series of characters that have ensured its establishment in NE Brazil. We focused on its reproductive biology and pollinator behavior to explain its reproductive success. We performed manual pollination and germination experiments, and monitored the behavior of C. retusa's main pollinators in monospecific plots, and in mixed plots where C. retusa occurred together with two congeners, Crotalaria pallida and Crotalaria lanceolata. Crotalaria retusa is self‐compatible and capable of automatic selfing. Inbreeding depression was expressed at the level of percent seed germination, but not seed set. Few insects visited the inflorescences. Legitimate pollinators were two large carpenter bees, Xylocopa frontalis and Xylocopa grisescens which, together, accounted for more than 90 percent of the visits. The former foraged on C. retusa exclusively and has low pollen spread potential. The latter flew longer distances between plants and visited fewer flowers per inflorescence, potentially increasing the extent of pollen carryover, but at the risk of increasing heterospecific pollen transfer, because it visits other Crotalaria species during the same foraging bout. The different foraging strategies, allied to morphological disadvantages represented by pollen overlap on X. grisescens' body, may partially explain the much lower seed germination observed in C. pallida and C. lanceolata. These results are consistent with the hypothesis that a reduction in flower constancy may significantly depress viable seed set by increasing the chances of self‐pollination.     

Pollinator effectiveness of native and non‐native flower visitors to an apparently generalist Andean shrub,Duranta mandonii (Verbenaceae)

Not all visitors to flowers are pollinators and pollinating taxa can vary greatly in their effectiveness. Using a combination of observations and experiments we compared the effectiveness of introduced honeybees with that of hummingbirds, native bees and moths on both the male and female components of fitness of the Andean shrub Duranta mandonii (Verbenaceae). Our results demonstrated significant variation among flower visitors in rates of visitation, pollen removal ability and contribution to fruit set. This variation was not always correlated; that is, taxa that regularly visited flowers did not remove the most pollen or contribute to fruit set. Despite the taxonomic diversity of visitors, the main natural pollinators of this shrub are large native bees, such as Bombus spp. Introduced honeybees were found to be as effective as native bees at pollinating this species. Duranta mandonii has high apparent generalization, but low realized generalization and can be considered to be a moderate ecological generalist (a number of species of large bees provide pollination services), but a functional specialist (most pollinators belong to a single functional group). The present study has highlighted the importance of measuring efficiency components when documenting plant–pollinator interactions, and has also demonstrated that visitation rates may give little insight into the relative importance of flower visitors.     

Models of floral pattern in detached flowers of Silene coeli-rosa (L) Godr. (Caryophyllaceae)

Organ number per whorl was analysed in aberrant flowers of the long-day (LD) plant , Silene coeli-rosa , to test a hypothesis that organ number in a whorl takes its cue from an adjacent outer whorl and that perturbed organ number per whorl is not random but defaults to that of closely related taxa or genera of the Caryophyllaceae. When plants were grown under short-days (SD), transferred to LD and the shoot meristem excised and cultured in vitro under SD, the normal pattern of flower development was often disrupted. For example, we observed flowers which comprised floral whorls with an aberrant number of floral organs. In part, this was an effect of tissue culture; however, the over-and-above effect was the establishment of an alternative pattern of development. Our data indicate that two distinct and recurrent patterns occurred in the aberrant flowers we observed in five separate experiments. First, pairs of floral whorls were linked so that aberration in one whorl resulted in the next whorl being more aberrant than normal. Second, the number of organs in aberrant whorls was not random, but defaulted to an organ number which mimicked the flowers of closely related species of Silene or related genera in the Caryophyllaceae.  © 2002 The Linnean Society of London , Botanical Journal of the Linnean Society , 2002, 140 , 229−235.     

Intraspecific divergence and convergence of floral tube length in specialized pollination interactions

Floral tubes are often thought to be a consequence of adaptive specialization towards pollinator morphology. We explore floral tube length evolution within Tritoniopsis revoluta (Iridaceae), a species with considerable geographical tube length variation. We ask whether tube lengths of T. revoluta populations are associated with pollinator proboscis lengths, whether floral divergence occurs in the presence of different pollinators and whether floral convergence occurs between distantly related populations pollinated by the same pollinator. Finally, we ask whether tube length evolution is directional. Shifts between morphologically different pollinators were always associated with shifts in floral morphology, even when populations were very closely related. Distantly related populations had similar tube lengths when they were pollinated by the same pollinator. Shifts in tube length tended to be from short to long, although reversals were not infrequent. After correcting for the population-level phylogeny, there was a strong positive, linear relationship between floral tube length and pollinator proboscis length, suggesting that plants are functionally specialized on different pollinators at different sites. However, because tube length evolution in this system can be a bidirectional process, specialization to the local pollinator fauna is unlikely to result in evolutionary or ecological dead-ends such as canalization or range limitation.     

展毛翠雀的花性状表型选择

传粉者的选择作用是花表型性状进化的重要驱动力, 解析选择作用的强度是理解花进化的关键。通过表型操控实验和表型选择研究能够分析花性状与其适合度的关系, 探究花性状的表型选择作用。为揭示花性状变化对雌性适合度的影响, 本研究处理展毛翠雀(Delphinium kamaonense var. glabrescens)花萼片大小, 并进行表型选择分析。结果表明: 人为减小展毛翠雀花萼片显著降低了传粉者的访花频率, 但是并没有影响种子产量(种子数和结籽率), 说明展毛翠雀花萼片的大小不影响种子产量, 可能主要吸引传粉昆虫输出花粉。通过雌性适合度(种子数量)估计表型选择梯度, 发现花萼片大小(长和宽)没有受到显著的直接选择梯度。但是, 花距长受到显著的线性选差和选择梯度, 表明花距的延长能够增加种子产量。本研究表明展毛翠雀花性状受到选择的作用, 但萼片和花距有不同的功能, 分别影响传粉者访问频率和种子产量。     

盗蜜对角蒿传粉者行为和生殖成功的影响

一些研究显示盗蜜对自交植物的结实和结籽没有显著影响。然而, 对于既有传粉者为其传粉实现异交又能通过自交实现生殖保障的兼性自交植物来说, 盗蜜对其生殖的影响还知之甚少。由于兼性自交植物可以自交, 盗蜜对其总体结实可能不会有显著影响, 但可能会通过影响传粉者行为而影响传粉者介导的结实。为了验证这一假说, 本研究以兼性自交的一年生角蒿(Invarvillea sinensis var. sinensis)为研究材料, 通过野外调查和控制实验, 探讨了盗蜜对传粉者介导的结实(传粉者行为)和总体结实率的影响。结果表明: 角蒿的盗蜜者和主要传粉者相同, 均为密林熊蜂(Bombus patagiatus)。熊蜂盗蜜频率平均为20.24% (范围为0-51.43%)。盗蜜对角蒿总体结实率、每果结籽数和每果种子重量没有显著影响。然而, 被盗蜜花的柱头闭合比率显著高于未被盗蜜花, 说明盗蜜影响传粉者的访花行为和传粉者介导的结实率。另外, 被盗蜜花的高度显著高于未被盗蜜花, 说明盗蜜者倾向于从较大较高的花上盗蜜。这些结果为全面认识盗蜜对植物生殖的影响提供了新的信息。     

Pollinators exert positive selection on flower size on urban,but not on rural Scotch broom (Cytisus scoparius L. Link)

Aims Adaptive evolution of invasive species is both particularly exciting for the evolutionary biologist and worrisome for those interested in controlling or halting spread. Invasive species often have a distinct timeline and well-recorded population expansion. As invaders encounter new environments, they undergo rapid adaptive evolution. Our aim in this study was to measure variation of floral size in the invasive shrub Cytisus scoparius (Scotch broom) and measure natural selection by pollinators on that trait. Past research has found that this invasive plant is pollinator limited in Washington State and that declines in pollinator populations can contribute to local extinction in another invaded range (New Zealand). This plant is pollinated by both native and introduced species of bees, representing a broad range of pollinator sizes. Cytisus scoparius has a flower structure that is highly conducive to studies on pollinator choice, even in the absence of direct pollinator observations.Methods We surveyed urban and rural sites in and around the city of Olympia in Washington State. Measuring banner width, we were able to show that flower size varies substantially between plants but minimally within plants. By measuring the proportion of flowers that were 'tripped', we could determine pollinator visitation rates and thus determine the level of selection due to pollinator choice alone.Important findings We found that C. scoparius is under natural selection by pollinators for increased flower size. However, such positive natural selection was only seen in urban populations although it was consistent across two flowering seasons. Rural populations of Scotch broom do not appear to be under selection on flower size. The natural selection by pollinators on broom flowers could result in adaptive evolution into a new pollination niche by an invading species. A higher level of variation in broom flowers seen here than seen in previous works in native regions suggests that C. scoparius may be highly diverse and primed for adaptive evolution.     

Flies and concealed nectar sources: morphological innovations in the proboscis of Bombyliidae (Diptera)

Bee-flies (Bombyliidae) have morphological adaptations of the mouthparts to particular floral traits. To investigate this the short, plesiomorphic proboscis of Hemipenthes morio was compared with the long, apomorphic proboscis of Bombylius major . A novel feeding position enables B. major to use flowers that open to the side as additional nectar sources. The new horizontal feeding position is enabled by the prolonged ventral base of the proboscis. Bombylius major exploits deep corolla tubes with an elongate proboscis, and an increased efficiency in both the suction pumps and the sealing mechanisms of the proboscis. The exploitation of narrow corolla tubes is made possible by the shift from a sponging feeding mode, exhibited by H. morio , to the exclusively sucking mode in B. major . Besides quantitative changes in the proportions of the different proboscis components, labellar movements as well as the structures of saliva distribution are changed along with this shift. The labial musculature of B. major does not significantly differ from the plesiomorphic state, since both examined species do not only feed on nectar, but also on pollen.     

Continuous characters in phylogenetic analyses: patterns of corolla tube length evolution in Lithospermum L. (Boraginaceae)

The present study comprises an analysis of six different scoring schemes and eight different types of analytic methods aiming to investigate the evolution of a continuous character (i.e. corolla tube length) in Lithospermum L. (Boraginaceae). Corolla tube length in the genus is quite variable, ranging from 1 mm to 75 mm, and the length of the corolla tube has implications for pollination biology, such as longer corolla tubes (> 25 mm in length) being pollinated by hummingbirds or moths. In general, the various methods resolve similar ancestral character states; however, different states are reconstructed at nodes in which the descendants greatly differ in corolla tube length. Additionally, it is suggested that all of the variation of a continuous character should be included in analyses, and this may necessitate multiple analyses with different partitions of the data. The various analyses provide evidence that two maximum parsimony methods, linear parsimony and the TNT method, minimize the number of different rates of evolution. In Lithospermum, six origins of corolla tubes > 20 mm in length are resolved, and these origins occurred at two different times periods: (1) in the shadow of hummingbird diversification in North America (approximately 6–8 Mya) and (2) more recently (approximately 1–1.5 MyA). Four substantial decreases in corolla tube length also are reconstructed, and these may be associated with the origin of self‐pollination. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

通泉草属3种植物柱头触敏性与花部特征之间的关系

选取3种具有不同传粉环境及花部结构差异明显的通泉草属(Mazus)植物,即毛果通泉草(M.spicatus Vaniot)、长蔓通泉草(M.longipes Bonati)和弹刀子菜(M.stachydifolius(Turcz.)Maxim.),观测其柱头触敏性行为,测量有关花部性状,对它们之间的相关性进行比较分析。结果显示,在相同强度的刺激下,毛果通泉草的柱头最为迟钝(闭合时间最长),弹刀子菜最为敏感,而长蔓通泉草介于两者之间。此外,柱头触敏性最强的弹刀子菜比柱头触敏性相对较弱的长蔓通泉草和毛果通泉草具有更深的花色、更大的单花展示、更长的药柱距以及更大的花粉胚珠比。研究结果表明通泉草属植物柱头触敏性的进化与花部特征密切相关,它们共同影响传粉过程中对传粉者的吸引、花粉落置、传递效率以及植物交配系统的构成式样。     

Phenotypic plasticity and integration in response to flooded conditions in natural accessions of Arabidopsis thaliana (L.) Heynh (Brassicaceae)

Flood response is a crucial component of the life strategy of many plants, but it is seldom studied in non-flooded tolerant species, even though they may be subjected to stressful environmental conditions. Phenotypic plasticity in reaction to environmental stress affects the whole plant phenotype and can alter the character correlations that constitute the phenotypic architecture of the individual, yet few studies have investigated the lability of phenotypic integration to water regime. Moreover, little has been done to date to quantify the sort of selective pressures that different components of a plant's phenotype may be experiencing under contrasting water regimes. Genetic differentiation and phenotypic plasticity at the single-trait and multivariate levels were investigated in 47 accessions of the weedy plant Arabidopsis thaliana, and the relationship of plastic characters to reproductive fitness was quantified. Results indicate that these plants tend to be highly genetically differentiated for all traits, in agreement with predictions made on the basis of environmental variation and mating system. Varied patterns of apparent selection under flooded and non-flooded conditions were also uncovered, suggesting trade-offs in allocation between roots and above-ground biomass, as well as between leaves and reproductive structures. While the major components of the plants' multivariate phenotypic architecture were not significantly affected by environmental changes, many of the details were different under flooded and non-flooded conditions.     

Functional differentiation in pollination processes among floral traits in Serapias species (Orchidaceae)

Floral displays, influencing attractiveness to insects, increase the number of pollinator visits and the efficiency of each visit in terms of pollen exchange and thus affect the plant reproductive success. Here, we conducted an in situ manipulation experiment to investigate whether the floral modifications affect reproductive success in natural orchid populations of Serapias lingua and Serapias vomeracea. We estimated male and female reproductive success of three treatment groups, disassembly of floral tube, cutting of lip, and painting of the callus surface, in terms of pollinaria removed/deposited and fruit production. Results revealed that phenotypic modification had opposite effects on reproductive success of two examine species. Indeed, reproductive success was significantly increased by the detached of the petals and sepals, and decreased, due to callus painting and lip removal, in S. lingua. On the contrary, unmanipulated plants of S. vomeracea showed significantly higher value of pollinaria removed and deposited and fruit set than manipulated ones. The differences between S. lingua and S. vomeracea agree to the different pollination strategy of examined species. S. vomeracea shows shelter imitation strategy, and thus, the disassembly of tunnel‐like corolla does not allow the insects to use the flower as a refuge, while S. lingua is a sexually deceptive orchid and therefore the opening of the flower made more visible callus (visible at a greater distance) increasing the pollinators attraction. This study provides evidence that pollinators were largely sensitive to the experimental modification of the flower phenotype, which is consistent with the presence of significant selection on individual floral characters. Our experimental investigations of the effects of variation in display on pollinator visitation provide insights into the evolution of floral morphology in orchid with shelter imitation strategy.     

Relationship between floral traits and floral visitors in two coexisting Tecoma species (Bignoniaceae)

In this paper, we studied the floral biology of Tecoma fulva ssp. garrocha and T.?stans in Vaqueros (Salta, Argentina), where both species coexist. We tested the idea that floral traits are associated with the pollinator types that visit them. According to our results, T.?fulva ssp. garrocha presented traits common to bird flowers and were visited by two species of Trochilidae (Colibri serrirostris and Chlorostilbon lucidus). In addition, T.?stans exhibited traits common to hymenoptera flowers and were visited principally by Apis mellifera and Bombus atratus. This study showed that floral traits are predictive of animal visitors.