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1.
The Orchidaceae characteristically contain a very large number of species that attract pollinators but do not offer them any form of reward in return for visitation. Such a strategy is highly unusual in the plant kingdom. We conducted experiments in order to manipulate the reward strategy of the rewardless bumble-bee-pollinated orchid Barlia robertiana by adding sucrose solution to inflorescences. We found that supplementation decreased the probability of a pollinator removing pollinia by approximately ten times. Despite pollinators visiting many more flowers per inflorescence on supplemented plants, eight times fewer pollinia were removed from supplemented inflorescences during each visit. Pollinia deposition patterns were not significantly affected by supplementation and no geitonogamous deposition was recorded. In populations where inflorescences were supplemented for 20 days, pollinia removal was reduced by over half for supplemented inflorescences, whereas fruit set was unmodified by supplementation. We conclude that rewardlessness would increase total seed paternity, but not change either total seed maternity or the probability that offspring were outcrossed in this species. To the authors' knowledge this is the first time that there has been an unequivocal experimental demonstration of an evolutionary advantage for rewardlessness in the Orchidaceae.  相似文献   

2.
European honeybees (Apis mellifera) were less efficient pollinators ofGrevillea barklyana than nectar-feeding birds. Nectar-collecting honeybees did not contact reproductive parts of flowers. Pollen-collecting honeybees preferentially visited malestage flowers but rarely visited female-stage flowers. Fruit set on caged inflorescences that allowed access to honeybees but excluded birds was reduced by more than 50% compared to inflorescences that were visited by both types of visitors. Further, fruit set on caged inflorescences was less than on bagged inflorescences that excluded both birds and honeybees, indicating that pollen removal by bees decreased opportunities for delayed autonomous selfing in the absence of birds. Although fruit set was not pollen-limited at the study site, pollen removal by honeybees would decrease fruit set in small populations where birds are scarce. In addition, pollen removal by honeybees would reduce opportunities for outcrossing and reproductive success through male function. Although honeybees have been in Australia for insufficient time to have exerted selection on floral traits, evolutionary shifts in response to these animals are likely to occur in the future.  相似文献   

3.
Abstract: Bougainvillea stipitata displays inflorescences with three pendant greenish flowers. Flowers open at sunset and last five days. Flower perianth is constricted in the middle and forms a tube that ends in five lobes. Fragrance is emitted by the papillae located on the lobe margins of the perianth. The nectary is located at the base of the staminal tube. It secretes fructose-dominant nectar with amino acids, phenols and reduced acids. Nectar secretion is continuous during flower lifetime and the flowers do not recover the reward. Mainly moths visit flowers. Pollen load on stigmas indicates that most flowers received more than 50 pollen grains, which are deposited by pollinators within the first two days of the flower life. Although the low natural fruit set of this self-incompatible species may be due to intraplant pollen flow, pollinators play an essential role for B. stipitata fruit production.  相似文献   

4.
Aims The hypothesis of predator satiation has been proposed to explain mast fruiting in various flowering plants. It considers that the simultaneous production of large numbers of seeds by a plant population reduces the risk of seed predation for each individual. Orchids produce huge numbers of seeds per fruit and rarely experience seed predation. It remains unclear which factors may affect fluctuating fruit production in orchids, which generally suffer a widespread pollen limitation. To explore the temporal pattern of fruiting and potential factors related to fluctuation in fruit production, we investigated reproductive success of a long-spurred orchid (Habenaria glaucifolia) in an alpine meadow with thousands of individuals over 8 years.Methods To estimate reproductive success, pollinator observation was conducted by day and at night, and pollinia removal and receipt were recorded in the field population for 8 years. To examine whether fruit set and seed set are pollen limited, we conducted supplementary pollination experiments and compared fruit set, seed set and pollinia movement of open-pollinated flowers from 2011 to 2013. We measured lengths of spurs and pollinator proboscises, and nectar volume and concentration, to identify potential pollinators.Important findings Hawkmoths were seen to be effective pollinators for H. glaucifolia in 3 years, whereas in the remaining 5 years no pollinators were observed, and consequently pollinia were rarely transferred. Numerous pollinia movements were observed in 2012, 2013 and 2014 (pollinia removal: 48, 59 and 85%; pollinia receipt 51, 70 and 80%), and correspondingly fruit set was significantly higher in 2012 and 2013 (59 and 46%) than in 2011 (25%). It was fruit set, rather than seed set, that was pollen limited in this orchid in the 3 years, in that supplementary pollination increased fruit set but did not increase seed set per fruit compared to natural. Three species of hawkmoths had proboscis lengths that matched the spur length of H. glaucifolia. Fruit set in this long spurred orchid depends on the activity of long-tongued hawkmoths, resulting in significant temporal variation in fruit production. Mast fruiting in this alpine orchid could be attributed to a 'sit and wait' strategy, awaiting an abundance of effective pollinators.  相似文献   

5.
Vertical raceme or spike inflorescences that are bee-pollinated tend to present their flowers horizontally. Horizontal presentation of flowers is hypothesized to enhance pollinator recognition and pollination precision, and it may also ensure greater consistency of pollinator movement on inflorescences. We tested the hypotheses using bee-pollinated Corydalis sheareri which has erect inflorescences consisting of flowers with horizontal orientation. We altered the orientation of individual flowers and prepared three types of inflorescences: (i) unmanipulated inflorescences with horizontal-facing flowers, (ii) inflorescences with flowers turned upward, and (iii) inflorescences with flowers turned downward. We compared number of inflorescences approached and visited, number of successive probes within an inflorescence, the direction percentage of vertical movement on inflorescences, efficiency of pollen removal and seed production per inflorescence. Deviation from horizontal orientation decreased both approaches and visits by leafcutter bees and bumble bees to inflorescences. Changes in floral orientation increased the proportion of downward movements by leafcutter bees and decreased the consistency of pollinator movement on inflorescences. In addition, pollen removal per visit and seed production per inflorescence also declined with changes of floral orientation. In conclusion, floral orientation seems more or less optimal as regards bee behavior and pollen transfer for Corydalis sheareri. A horizontal orientation may be under selection of pollinators and co-adapt with other aspects of the inflorescence and floral traits.  相似文献   

6.
Keasar T 《Animal behaviour》2000,60(5):639-646
Many species of orchids that do not offer food rewards to pollinators bloom in clusters, early in the season, and are polymorphic for corolla colour. Previous studies suggest that the foraging behaviour of insect pollinators may select for early blooming and colour polymorphism. I tested whether pollinator behaviour can also favour aggregated flowering in these species, in a two-stage laboratory experiment on na?ve bumblebees, Bombus terrestris (L.). In the first stage, the bees were allowed to forage on three colours of artificial flowers that contained sucrose rewards. In the second stage, I added nonrewarding flowers of a fourth colour and recorded the bees' visits to them. The four types of artificial flowers were either arranged in spatially distinct clusters, or were randomly intermingled. I used two reward schedules for each spatial arrangement: constant refilling of reward-containing flowers and probabilistic refilling. Bees that foraged on clustered flowers flew more often to the nonrewarding patch, and made more visits to nonrewarding flowers, than bees that foraged on intermingled flowers. This tendency was obtained both in the constant reward and in the probabilistic reward schedules. The results support the hypothesis that pollinator attraction may select for clustered, synchronized blooming in flowers that do not contain nectar and pollen rewards. Copyright 2000 The Association for the Study of Animal Behaviour.  相似文献   

7.
Morning floral heat as a reward to the pollinators of the Oncocyclus irises   总被引:2,自引:0,他引:2  
Sapir Y  Shmida A  Ne'eman G 《Oecologia》2006,147(1):53-59
Relationships between flowering plants and their pollinators are usually affected by the amount of reward, mainly pollen or nectar, offered to pollinators by flowers, with these amounts usually positively correlated with floral display. The large Oncocyclus iris flowers, despite being the largest flowers in the East Mediterranean flora, are nectarless and have hidden pollen. No pollinators visit the flowers during daytime, and these flowers are pollinated only by night-sheltering solitary male bees. These iris flowers are partially or fully dark-colored, suggesting that they gather heat by absorbing solar radiation. Here we test the hypothesis that the dark-colored flowers of the Oncocyclus irises offer heat reward to their male solitary bee pollinators. Floral temperature was higher by 2.5°C than ambient air after sunrise. Solitary male bees emerged earlier after sheltering in Oncocyclus flowers than from other experimental shelter types. Pollination tunnels facing east towards the rising sun hosted more male bees than other aspects. We suggest that floral heat reward can explain the evolution of dark floral colors in Oncocyclus irises, mediated by the pollinators’ behavior.  相似文献   

8.
Summary Bumble bees (Bombus spp.) picked up a pollen packet (pollinium) of milkweed (Asclepias syriaca L.) every 2–5 h, usually more rapidly on their feet than on their mouthparts. Pollinia were retained an average of slightly over one day on the mouthparts and one-quarter day on the feet. This long retention period enhances the possibility of outcrossing in this largely or completely self-incompatible species. Although many more pollinia were carried on the feet, the longer retention of those on the mouthparts resulted in their collective outcrossing potential exceeding that of pollinia on the feet. Once pollinia became attached to the bees (via their attachment mechanisms, the corpuscula), others often attached to the translator arms (connecting corpusculum with pollinia) of the first pollinia. Long strings of pollinia and corpuscula often resulted, but they frequently were shed down to a single corpusculum, which was usually retained for long periods (estimated retention time=12–43 days). During the middle and latter part of the flowering period about two-thirds of the bees' feet carried only a single corpusculum. Feet in this condition picked up extremely few pollinia and thus were largely unavailable as sites for pollen transfer. Having several single corpuscula on the feet greatly lowered the number of pollinia carried by a bee over the season. Single corpuscula occurred much less frequently on the mouthparts and were shed over 25 times as rapidly as those on the feet. Many more bumble bees moved between clones than did other possible diurnal pollinators, largely a result of being several times more abundant than all other visitors combined. This factor, plus their heavy pollinia load, suggests that they were the most important diurnal pollinators of these clones.  相似文献   

9.
Deceptive orchid species that offer no floral rewards to pollinators often experience pollinator limitation because they. The breeding system and fruit set were investigated in order to examine pollinator limitation for a population of a deceptive orchid, Pogonia japonica , on a floating peat mat in Mizorogaike Pond, western Japan. Fruit sets for outcross-pollinated, self-pollinated and pollinator-excluded flowers were 75%, 80%, and 0%, respectively. Thus, this species was self-compatible but neither autogamous nor apogamous. The fruit set for open-pollinated flowers was 22.9% and 17.4% in 1994 and 1995, suggesting that pollinator limitation did occur in the field. Larger flowers were more likely to develop into fruit than smaller ones in open pollination (36% vs 8%). Thus, flower size seems to affect fruiting of this species. No effective visits by major pollinators such as bees, which could carry pollinia, were observed during 16.5 h of observation during daytime. Instead, thrips were often found moving on gynostemium or pollinia and circumstantial evidence that suggests thrips may partly transfer pollen in pieces out of a granular pollinium of P. japonica was obtained. Pogonia japonica should come to use thrips as a pollinator to supplement low fruit set under limited flower visitation by major pollinators.  相似文献   

10.
The inflorescences of Phyla incisa consist of flowers in two phases: younger, nectar-containing flowers that have yellow corolla throats and older, nectar-lacking flowers that have dark purple corolla throats. Observations of pollinator visitation patterns to both natural and manipulated inflorescences were made to determine the role of each flower phase in pollinator attraction. The effect of older-phase flowers on male and female reproductive success was determined by comparing stigmatic pollen loads and estimates of pollen removal from inflorescences having different numbers of these flowers. The pollinators of Phyla selected larger inflorescences more often than expected based upon the size distribution of inflorescences available to them. Both younger- and older-phase flowers contributed to the attraction of pollinators, but the latter were less effective in this function. The presence of older-phase flowers significantly increased the visitation rate to inflorescences and the amount of pollen removed but had little effect on pollen deposition on stigmas. The lack of correspondence between pollen deposition and pollinator-visitation rate was not due to stigma saturation, since stigma loads varied greatly. The data indicate that the deposition of pollen on stigmas in this species is a relatively stochastic process, whereas pollen removal from inflorescences occurs at a much more regular rate. Old-phase flower retention appears to contribute to reproductive success through increased pollen donation when pollinator activity is high and may also increase the probability of seed set when pollinators are rare.  相似文献   

11.
Thelymitra epipactoides has a highly variable visual display achieved through polychromatic flowers and variable inflorescence size, bearing between 7 and 31 flowers, which attract foraging polylectic bees. Only bees of the genusNomia were observed carrying pollinia and successfully pollinating the orchid. The genusNomia contains polylectic, pollen gathering species that store pollen in both the crop and scopa on the hind legs. The absence of a reward for the bees indicates the orchid is relying on deception to attract visitors. The relationship of deception to mimicry is discussed. Once on the flower, tactile, visual and possibly olfactory stimuli direct bees to the false anther formed by the voluminous column wings, where morphological adaptations of the flower ensure that the pollinarium is deposited on the gaster of the bee to effect pollination. — The lack of seed set observed on the Victorian coast appears to be due to the absence of pollinators from the heath and grassland communities in which the orchid grows. This may well be a consequence of the reduced number of plants flowering in the community (a result of the elimination of fire at these sites), thus not maintaining a floral community attractive to potential pollinators.  相似文献   

12.
Summary Although it is known that stigmatic pollen deposition may trigger early flower senescence, the existence of a similar plastic response of flower lifespan to pollen removal has been much less studied. Here we report on a factorial, manipulative experiment in which all 2 × 2 flower combinations of pollinia removal and stigmatic pollinia insertion were performed in inflorescences of the Patagonian ground orchidChloraea alpina. This experiment was conducted in the laboratory, in a population of cut inflorescences and in the field. We hypothesized that if expected fitness gains, through both the male and female functions, were weighed against the costs of flower maintenance, then early flower senescence should be triggered by either pollinia removal or insertion. The shortest flower lifespan would be expected in flowers where both processes occurred. Results showed that flower longevity was very strongly affected by pollinia insertion, reducing the flower lifespan by approximately 60%. The response of pollinia removal was much weaker. A significant reduction in flower longevity caused by pollinia removal was only detected in unpollinated flowers (i.e. no pollinia inserted). Within the racemose inflorescences, flowers in basal positions lived longer than flowers in terminal ones, which might be evidence of the importance of resource availability in determining maximum flower longevity. The observed responses of flower lifespan plasticity to pollinia manipulation only partially supported our expectations based on fitness benefit—cost relationships. Other factors that might explain these discrepancies are the different fitness gains that may indeed accrue to the processes of pollinia removal and insertion as they occur in nature, donor manipulation of the recipient flower lifespan associated with the evolution of pollen clustering into pollinia and physiological constraints in terms of the extent to which flower longevity may respond to pollen removal.  相似文献   

13.
The floral visitors of silky oak, Grevillea robusta A. Cunn. ex R.Br., their foraging behaviour and their effects on fruit‐set were studied at Malava, western Kenya. Grevillea robusta is a popular tree for farm plantings in the eastern and central African highlands. Yield of seed has been disappointingly low in some areas and a lack of appropriate pollinators has been suggested as a possible cause. Investigations involved the monitoring of visitors on active inflorescences, assessment of the rewards available to potential pollinators, and exclusion experiments to establish the effects of various visitors on fruit‐set. The flowers are visited mainly by birds and insects. The likely pollinators of G. robusta are sunbirds (Nectarinia amethystina, N. cyanolaema, N. olivacea, N. superba and N. venusta) and white‐eyes (Zosterops kikuyuensis and Z. senegalensis). Very little aggressive behaviour between birds was recorded. No nocturnal pollinators were observed. Nectar was the major floral reward for pollinators, but is likely depleted by ants and honey bees, the foraging behaviour of which confirmed them to be nectar‐robbers. These insects hardly ever touched stigmas during their visits. Eighty‐nine per cent of bird visits were in the morning (07.00–10.00 hours) when nectar volume was highest. Inflorescences bagged to exclude birds set no fruits, and unmanipulated flowers and flowers bagged with self‐pollen set no fruits, indicating a self‐incompatibility mechanism. Control cross‐pollinated flowers displayed greatly increased fruit‐set (25.1%) compared with natural open‐pollination (0.9%). All these findings confirm the importance of cross‐pollen transfer to flowers and the necessity of pollinators for fruit‐set. Effective seed production requires activity of pollinators for self‐pollen removal and cross‐pollen deposition. Seed production stands for G. robusta should be established where flowering is prolific and bird pollinators are abundant.  相似文献   

14.
Most hermaphroditic, many-flowered plants should suffer reduced fitness from within-plant selfing (geitonogamy). Large inflorescences are most attractive to pollinators, but also promote many flower visits during a single plant visit, which may increase selfing and decrease pollen export. A plant might avoid the negative consequences of attractiveness through modification of the floral display to promote fewer flower visits, while retaining attractiveness. This report shows that increasing only the variance in nectar volume per flower results in fewer flower visits per inflorescence by wild hummingbirds ( Selasphorus rufus ) and captive bumble bees ( Bombus flavifrons ) foraging on artificial inflorescences. Inflorescences were either constant (all flowers contained the same nectar volume) or variable (half the flowers were empty, the other half contained twice as much nectar as in the constant flowers). Both types of inflorescence were simultaneously available to foragers. Risk-averse foraging behaviour was expressed as a patch departure preference: birds and bees visited fewer flowers on variable inflorescences, and this preference was expressed when resource variability could be determined only by concurrent sampling. When variance treatments were clearly labelled with colour and offered to hummingbirds, the departure effect was maintained; however, when preference was measured by inflorescence choice, birds did not consistently prefer to visit constant inflorescences. The reduced visitation lengths on variable inflorescences by both birds and bees documented in this study imply that variance in nectar production rates within inflorescences may represent an adaptive trait to avoid the costs of geitonogamy.  相似文献   

15.
Summary Bumblebees foraging on vertical inflorescences start near the bottom and work upward, behavior commonly interpreted as a response to the greater amounts of nectar available in lower flowers. Lupinus polyphyllus, which produces no nectar, has more pollen available in upper flowers. Although bees are probably unable to detect this gradient, since pollen is hidden from their view, they still start low and forage upward. Therefore, we concluded that the bees' tendency to forage upward on vertical inflorescences is not tied to a reward gradient. In addition, bees use only about 15% of the flowers per inflorescence, although they could be much more efficient by visiting and revisiting every flower systematically. In general, revisits would not be penalized because most flowers contain enough pollen for several visits. Optimal foraging theory may not offer an adequate explanation for such gross inefficiency.  相似文献   

16.
Pollinators visit flowers for rewards and should therefore have a preference for floral signals that indicate reward status, so called ‘honest signals’. We investigated honest signalling in Brassica rapa L. and its relevance for the attraction of a generalised pollinator, the bumble bee Bombus terrestris (L.). We found a positive association between reward amount (nectar sugar and pollen) and the floral scent compound phenylacetaldehyde. Bumble bees developed a preference for phenylacetaldehyde over other scent compounds after foraging on B. rapa. When foraging on artificial flowers scented with synthetic volatiles, bumble bees developed a preference for those specific compounds that honestly indicated reward status. These results show that the honesty of floral signals can play a key role in their attractiveness to pollinators. In plants, a genetic constraint, resource limitation in reward and signal production, and sanctions against cheaters may contribute to the evolution and maintenance of honest signalling.  相似文献   

17.
A great diversity of flower morphology in orchids has long been thought to be selected by diverse pollinators. Habenaria Willd. (Orchidaceae) species are generally characterized by long nectar spurs and pollinated by long‐tongued insects (Lepidoptera), the mechanical fit between the spur and pollinator proboscis length being supposedly caused by “arms race” reciprocal selection. Here, we report that flowers of Habenaria aitchisonii Rchb. f. with nectar spurs (approximately 9 mm) were pollinated by three species of settling noctuid moths whose proboscises varied in length from 10 to 16 mm. When a settling moth crawled on the spikes and probed the flowers for nectar, pollinia were placed on the moths’ legs rather than on other body parts. Our 5‐year survey of pollinia movement and 3‐year supplemental pollination experiments indicated that fruit and seed production in this orchid were not often pollen‐limited at flower level. In a natural population in Shangri‐La, Southwest China, the proportions of pollinia removal and deposition on stigmas by moth legs were 93.8% and 83.5%, respectively. This finding of efficient pollen transfer by the pollinators’ legs in H. aitchisonii adds a new example of diverse pollinia placement on pollinators (here settling moths) in the Orchidaceae.  相似文献   

18.
Many orchid species are unusual in that they provide no nectar or pollen rewards for their pollinators. Absence of reward is expected to have a fundamental effect on pollinator visitation patterns. In particular the number of flowers visited per inflorescence is expected to be affected in both unrewarding and co-flowering rewarding species. We used arrays of artificial inflorescences, which could be either rewarding or unrewarding and were differentiated by their colour, to test how many flowers bumblebees visit in each type of inflorescence. The frequency of the two colours was varied, thus modelling the case where different frequencies of both an unrewarding and rewarding species were present in a patch. We found that bumblebees visited more flowers per rewarding inflorescence after they have experienced unrewarding or partially emptied rewarding inflorescences. We used these results to simulate pollen transfer and thus predict selfing rates on rewarding inflorescences. We found these increased when nectar depleted or when there was a greater proportion of unrewarding inflorescences in the patch. Conversely, we found that the number of flowers bumblebees visited on each unrewarding inflorescence did not significantly change through experiments. Selfing rates for unrewarding inflorescences were predicted to depend principally on the number of these inflorescences bumblebees visited rather than on the number of flowers they visit per inflorescence. This was because most visitors to orchids are supposed to be naive, and pollinators that commence foraging carrying no pollen will necessarily self any flower they pollinate on the first inflorescence they visit. Thus the average selfing rate is expected to increase as the sequence of inflorescences visited decreases in length.  相似文献   

19.
 The terrestrial orchid Epipactis helleborine is a morphologically variable species with a wide distribution in Europe. It is pollinated by social wasps, and most populations show the morphological characteristics of outcrossing species. However, local predominantly selfing subspecies and varieties have been documented from drier habitats. To document geographic variation in floral morphology, ability to produce seeds through autogamy, and reproductive success in E. helleborine, we sampled 13 populations from three geographic regions along a latitudinal gradient of c. 1000 km from northern to southern Sweden. In the southernmost region, populations in dry and mesic habitats were compared. Supplemental hand-pollination was conducted to determine whether among-population variation in fruit set could be explained by differences in the natural level of pollination, and whether any relationship between floral morphology and fruit production could be explained by interactions with pollinators. Bagging experiments showed no evidence of autogamy in any of the study populations. Number of flowers, pollinia removal and fruit set varied significantly among populations but did not differ among regions. Pollinia removal was positively correlated with population size and both pollinia removal and fruit set were lower in dry than in mesic habitats. At the level of the individual plant, the number of pollinia removed increased more rapidly with flower number than did number of fruits produced. The hand-pollination experiment indicated that the positive relationship between number of flowers and fruit production was due to a higher degree of pollen limitation in plants with few flowers than in plants with many flowers. The experiment also showed that variation in the level of pollen limitation could only partly explain variation in fruit set among populations. Received November 6, 2001; accepted April 27, 2002 Published online: December 3, 2002  相似文献   

20.
Low fruit set values in most orchids (especially epiphytic and tropical species) are normally thought to be the consequence of pollination constraints and limited resources. In particular, pollination constraints are modulated by pollinator visitation rates, pollinator visitation behaviour (promoting crossing or selfing), the type and number of pollinia deposited on stigmas (in the case of orchids with subequal pollinia) and the amount of pollen loaded per inflorescence. In order to assess to what extent these factors can affect fruit set in specific orchid-pollinator systems, the repercussions of some of these aspects on reproduction of Broughtonia lindenii were examined in a coastal population in western Cuba. The study focused on plant breeding system, importance of pollen load and type of pollinia on subsequent fruit and seed, limiting factors of seed production and interaction with pollinators. This species presents long-lasting flowers that senesce after all forms of effective visit. Pollinator dependence for fruit production was demonstrated, while hand-pollination experiments revealed self-compatibility and inbreeding depression at seed level. More pollinia on stigmas enhance the proportion of well-developed seeds. In contrast, the pollinia type used in pollination is not important for seed quality of fruits, suggesting that small pollinia are not rudimentary. Natural fruit set in two consecutive years was substantially affected by pollinator activity, and also by systematic depredatory activity of ants and a caterpillar. Considering that this orchid completely lacks nectar and that the local assemblage of pollinators and predators influenced its reproduction, a minor importance of resource constraints in this epiphyte (with long-lasting reserve structures) is confirmed at least for a short time.  相似文献   

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