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1.

The pollination syndrome concept implies that flowers evolved with particular sets of characteristics, such as colors, shapes, orientations, and rewards, as a means of attracting pollinators. However, these traits may have also evolved to deter unwanted visitors. The North American genus Penstemon exhibits a great floral diversity that is mainly associated with bumblebee and hummingbird pollination. Evolutionary shifts from insect pollination to hummingbird pollination have occurred in Penstemon repeatedly, but some species maintain mixed-pollination systems and intermediate floral traits between bee- and hummingbird-pollination modes. The apparently intermediate floral traits of species with mixed-pollination systems might be potentially acting to deter bumblebee foragers. Then, bird-flower traits might be selected with increased hummingbird visitation over evolutionary time might, resulting in specialization to and the evolution of floral traits present in hummingbird-pollinated species. Here, we modified bee-pollination floral traits in Penstemon gentianoides with a mixed pollination system, to resemble hummingbird-pollination traits, and measured the effects of trait modification on bumblebee foraging behavior and plant female reproductive fitness. Our results showed that reduction in the width of the corolla tube and the absence of the corolla lip negatively affects bumblebee visitation and their efficiency as pollinators, and that the synergistic interaction of both traits enhanced the “anti-bee” effect. We conclude that acquisition of floral traits that resemble those of hummingbird-pollination enables Penstemon plant species to deter bumblebee visits.

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2.
Long-tubed hawkmoth-pollinated species present some of the most remarkable examples of floral specialization depending exclusively on long-tongued hawkmoths for sexual reproduction. Nonetheless, long-tongued hawkmoths do not rely exclusively on specialized plants as nectar sources, which may limit sexual reproduction through pollen limitation. However, very few studies have quantified the level of pollen limitation in plants with highly specialized floral traits in tropical regions. In this context, we studied four sympatric hawkmoth-pollinated species in a highland Atlantic Rain forest and assessed pollen limitation and their dependence on pollinators by analyzing the floral biology, breeding system, pollination mechanisms, and abundance of long-tongued pollinators. We showed that the four species are self-compatible, but are completely dependent on long-tongued hawkmoths to set fruits, and that flower visitation was infrequent in all plant species. Pollen limitation indices ranged from 0.53 to 0.96 showing that fruit set is highly limited by pollen receipt. Long-tongued moths are much less abundant and comprise only one sixth of the hawkmoth fauna. Pollen analyses of 578 sampled moths revealed that hawkmoths visited ca. 80 plant species in the community, but only two of the four species studied. Visited plants included a long-tubed hawkmoth-pollinated species endemic to the lowland forest ca. 15–20 km away from the study site. Specialization index (H 2 ?=?0.20) showed that community-level interactions between hawkmoths and plants are generalized. We suggest that sexual reproduction of these highly specialized hawkmoth-pollinated species is impaired by competition among plants for pollinators, in conjunction with the low abundance and diversity of long-tongued pollinators.  相似文献   

3.
The Canary Islands are home to a guild of endemic, threatened bird-pollinated plants. Previous work has suggested that these plants evolved floral traits as adaptations to pollination by flower specialist sunbirds, but subsequently, they appear to have co-opted generalist passerine birds as sub-optimal pollinators. To test this idea, we carried out a quantitative study of the pollination biology of three of the bird-pollinated plants, Canarina canariensis (Campanulaceae), Isoplexis canariensis (Veronicaceae) and Lotus berthelotii (Fabaceae), on the island of Tenerife. Using colour vision models, we predicted the detectability of flowers to bird and bee pollinators. We measured pollinator visitation rates, nectar standing crops as well as seed-set and pollen removal and deposition. These data showed that the plants are effectively pollinated by non-flower specialist passerine birds that only occasionally visit flowers. The large nectar standing crops and extended flower longevities (>10 days) of Canarina and Isoplexis suggests that they have evolved a bird pollination system that effectively exploits these low frequency non-specialist pollen vectors and is in no way sub-optimal. Seed set in two of the three species was high and was significantly reduced or zero in flowers where pollinator access was restricted. In L. berthelotii, however, no fruit set was observed, probably because the plants were self-incompatible horticultural clones of a single genet. We also show that, while all three species are easily detectable for birds, the orange Canarina and the red Lotus (but less so the yellow-orange Isoplexis) should be difficult to detect for insect pollinators without specialised red receptors, such as bumblebees. Contrary to expectations if we accept that the flowers are primarily adapted to sunbird pollination, the chiffchaff (Phylloscopus canariensis) was an effective pollinator of these species.  相似文献   

4.
Both plant traits and plant–pollinator interactions are thought to influence plant mating systems. For hummingbird-pollinated plants, foraging strategy (territorial or traplining) is also expected to influence plant mating. We hypothesize that the traplining behavior of hermits promotes outcrossing, whereas the behavior of non-hermits favours self-incompatibility. Thus, selection is expected to maintain self-incompatibility in plants pollinated by non-hermits. We explore the incidence of self-incompatibility in Neotropical hummingbird-pollinated plants and its association with hummingbird behavior and plant traits. We conducted a literature review (56 species) and performed hand-pollination experiments in 27 hummingbird-pollinated plants in an Atlantic rainforest. We found that self-incompatibility (measured as <0.3 for the Index of Self-incompatibility [ISI]) occurred in only 33 % of the Neotropical hummingbird-pollinated plants. The interaction of hummingbird and habit type affected ISI, as did phylogenetic relationships. Specifically, herbs pollinated by non-hermits had higher ISI than woody plants pollinated by non-hermits, and herbs pollinated by both hermits and non-hermits. For the Atlantic rainforest plant guild, 30 % of the species were self-incompatible. ISI was higher in herbs than in woody species and increased with plant aggregation but was not dependent on foraging behavior, plant density, or floral display. Although hummingbirds differ in their foraging strategies, these behavioral differences seem to have only a minor influence on the incidence of self-incompatibility. Phylogenetic relatedness seems to be the strongest determinant of mating system in Neotropical hummingbird-pollinated plants.  相似文献   

5.
In generalised pollination systems, the presence of alien plant species may change the foraging behaviour of pollinators on native plant species, which could result in reduced reproductive success of native plant species. We tested this idea of indirect interactions on a small spatial and temporal scale in a field study in Mauritius, where the invasive strawberry guava, Psidium cattleianum, provides additional floral resources for insect pollinators. We predicted that the presence of flowering guava would indirectly and negatively affect the reproductive success of the endemic plant Bertiera zaluzania, which has similar flowers, by diverting shared pollinators. We removed P. cattleianum flowers within a 5-m radius from around half the B. zaluzania target plants (treatment) and left P. cattleianum flowers intact around the other half (control). By far, the most abundant and shared pollinator was the introduced honey bee, Apis mellifera, but its visitation rates to treatment and control plants were similar. Likewise, fruit and seed set and fruit size and weight of B. zaluzania were not influenced by the presence of P. cattleianum flowers. Although other studies have shown small-scale effects of alien plant species on neighbouring natives, we found no evidence for such negative indirect interactions in our system. The dominance of introduced, established A. mellifera indicates their replacement of native insect flower visitors and their function as pollinators of native plant species. However, the pollination effectiveness of A. mellifera in comparison to native pollinators is unknown. Christine B. Müller, deceased 7 March 2008.  相似文献   

6.
Following the recent discovery of rodent pollination in the Pagoda lily, Whiteheadia bifolia (Hyacinthaceae) in South Africa, now the Cape rock elephant-shrew, Elephantulus edwardii (Macroscelidea, Afrotheria) is reported as an additional pollinator. Elephant-shrews, live-trapped near W. bifolia plants, were released in two terraria, containing the plants. The animals licked nectar with their long and slender tongues while being dusted with pollen and touching the stigmas of the flowers with their long and flexible noses. The captured elephant-shrews had W. bifolia pollen in their faeces, likely as a result of grooming their fur as they visited the flowers without eating or destroying them. The animals mostly preferred nectar over other food. This is the first record of pollination and nectar consumption in the primarily insectivorous E. edwardii, contributing to the very sparse knowledge about the behaviour of this unique clade of African mammals, as well as pollination by small mammals.  相似文献   

7.
Pollinators search for multiple flora resources throughout their life cycle. Most studies, however, only assess how bees discriminate floral cues in the context of nectar foraging. In the present study, we sought to elucidate whether oil-collecting bees discriminate flowers of Byrsonima variabilis (Malpighiaceae) with petals of different colours when foraging for pollen or oil. As the colour of the standard petal changes during anthesis, we characterised the spectral reflectance patterns of flowers throughout anthesis and modelled chromatic perceptual space to determine how these colour patterns are perceived by bees. Through the quantification of flower pollen in the different phases, we found that the colour of the standard petal is an honest cue of the presence of pollen. Centridine bees preferentially visited flowers with a yellow (bee’s green) colour when searching for pollen, but indiscriminately visited flowers with different petal colours when searching for floral oil. We suggest that standard petals, in the species studied and others of the genus, like nectar guides, act as pollen guides, which oil-collecting females use to detect pollen-rich flowers. Moreover, they use different floral clues during foraging for different resources in the same host plant.  相似文献   

8.
Plants offer metabolically rich floral nectar to attract visiting pollinators. The composition of nectar includes not only sugars, but also amino acids. We have examined the amino acid content of the nectar of ornamental tobacco and found that it is extremely rich (2 mM) in proline. Because insect pollinators preferentially utilize proline during the initial phases of insect flight and can reportedly taste proline, we determined whether honeybees showed a preference for synthetic nectars rich in proline. We therefore established an insect preference test and found that honeybees indeed prefer nectars rich in the amino acid proline. To determine whether this was a general phenomenon, we also examined the nectars of two insect-pollinated wild perennial species of soybean. These species also showed high levels of proline in their nectars demonstrating that plants often produce proline-rich floral nectar. Because insects such as honeybees prefer proline-rich nectars, we hypothesize that some plants offer proline-rich nectars as a mechanism to attract visiting pollinators.  相似文献   

9.
Flower colour is an important signal used by flowering plants to attract pollinators. Many anthophilous insects have an innate colour preference that is displayed during their first foraging bouts and which could help them locate their first nectar reward. Nevertheless, learning capabilities allow insects to switch their colour preferences with experience and thus, to track variation in floral nectar availability. Manduca sexta, a crepuscular hawkmoth widely studied as a model system for sensory physiology and behaviour, visits mostly white, night-blooming flowers lacking UV reflectance throughout its range in the Americas. Nevertheless, the spectral sensitivity of the feeding behaviour of naïve moths shows a narrow peak around 450 nm wavelengths, suggesting an innate preference for the colour blue. Under more natural conditions (i.e. broader wavelength reflectance) than in previous studies, we used dual choice experiments with blue- and white-coloured feeders to investigate the innate preference of naïve moths and trained different groups to each colour to evaluate their learning capabilities. We confirmed the innate preference of M. sexta for blue and found that these moths were able to switch colour preferences after training experience. These results unequivocally demonstrate that M. sexta moths innately prefer blue when presented against white flower models and offer novel experimental evidence supporting the hypothesis that learning capabilities could be involved in their foraging preferences, including their widely observed attraction to white flowers in nature.  相似文献   

10.
Floral volatile organic compounds (VOCs) play important roles in plant-pollinator interactions. We investigated the reproductive ecology and floral VOCs of Zygopetalinae orchids to understand the relationship between floral scents and pollinators. We performed focal observations, phenological censuses and breeding system experiments in eight species in southeast Brazil. Floral scents were collected and analysed using SPME/GC-MS. We performed multivariate analyses to group species according to affinities of their VOCs and define compounds associated to each plant. Dichaea cogniauxiana was pollinated by weevils which use their developing ovules, while D. pendula was pollinated by the same weevils and perfume-collecting male euglossine bees. The other species were deceit-pollinated by bees. Zygopetalum crinitum was pollinated by carpenter bees, while W. warreana, Z. mackayi and Z. maxillare were bumblebee-pollinated. The latter was also pollinated by Centris confusa. Breeding system varied widely with no association to any pollinator group. Most VOCs are common to other floral scents. Zygopetalum crinitum presented an exclusive blend of VOCs, mainly composed of benzenoids. The scents of Pabstia jugosa, Promenaea xanthina and the Zygopetalum spp. were similar. The bumblebee-pollinated species have flowering periods partially overlapped, thus neither phenology nor pollinators constitute hybridization barriers among these species. Euglossines are not the only pollinators of Zygopetalinae. Different VOCs, size and lifespan of flowers are associated with distinct pollinators. A distinctive VOC bouquet may determine specialisation in carpenter bees or male euglossines within bee-pollinated flowers. Finally, visitation of deceit-pollinated flowers by perfume-collecting euglossines allows us to hypothesise how pollination by this group of bees had evolved.  相似文献   

11.
As part of a study of the roles of the sensory subsystem devoted to CO2 in the nectar-feeding moth Manduca sexta, we investigated CO2 release and nectar secretion by flowers of Datura wrightii, a preferred hostplant of Manduca. Datura flowers open at dusk and wilt by the following noon. During the first hours after dusk, when Manduca feeds, the flowers produce considerable amounts of nectar and emit levels of CO2 that should be detectable by moths nearby. By midnight, however, both nectar secretion and CO2 release decrease significantly. Because nectar production requires high metabolic activity, high floral CO2 emission may indicate food abundance to the moths. We suggest that hovering moths could use the florally emitted CO2 to help them assess the nectar content before attempting to feed in order to improve their foraging efficiency.Electronic Supplementary Material  Supplementary material is available in the online version of this article at  相似文献   

12.
Flowers adapted for hummingbird pollination are typically red. This correlation is usually explained by the assertion that nectar- or pollen-stealing bees are “blind” to red flowers. However, laboratory studies have shown that bees are capable of locating artificial red flowers and often show no innate preference for blue over red. We hypothesised that these findings might be artefacts of the simplified laboratory environment. Using bumblebees (Bombus impatiens) that had been trained to visit red and blue artificial flowers, we tested whether colour preference was influenced by complexity of the background on which they were foraging. Many bees were indifferent to flower colour when tested using a uniform green background like those commonly used in laboratory studies, but all bees showed strong colour preferences (usually for blue) when flowers were presented against a photograph of real foliage. Overall, preference for blue flowers was significantly greater on the more realistic, complex background. These results support the notion that the red of “hummingbird syndrome” flowers can function to reduce bee visits despite the ability of bees to detect red and highlight the need to consider context when drawing inferences about pollinator preferences from laboratory data. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.  相似文献   

13.
On the success of a swindle: pollination by deception in orchids   总被引:1,自引:0,他引:1  
A standing enigma in pollination ecology is the evolution of pollinator attraction without offering reward in about one third of all orchid species. Here I review concepts of pollination by deception, and in particular recent findings in the pollination syndromes of food deception and sexual deception in orchids. Deceptive orchids mimic floral signals of rewarding plants (food deception) or mating signals of receptive females (sexual deception) to attract pollen vectors. In some food deceptive orchids, similarities in the spectral reflectance visible to the pollinator in a model plant and its mimic, and increased reproductive success of the mimic in the presence of the model have been demonstrated. Other species do not mimic specific model plants but attract pollinators with general attractive floral signals. In sexually deceptive orchids, floral odor is the key trait for pollinator attraction, and behaviorally active compounds in the orchids are identical to the sex pheromone of the pollinator species. Deceptive orchids often show high variability in floral signals, which may be maintained by negative frequency-dependent selection, since pollinators can learn and subsequently avoid common deceptive morphs more quickly than rare ones. The evolution of obligate deception in orchids seems paradoxical in the light of the typically lower fruit set than in rewarding species. Pollination by deception, however, can reduce self-pollination and encourage pollen flow over longer distances, thus promoting outbreeding. Although some food deceptive orchids are isolated through postzygotic reproductive barriers, sexually deceptive orchids lack post-mating barriers and species isolation is achieved via specific pollinator attraction. Recent population genetic and phylogenetic investigations suggest gene-flow within subgeneric clades, but pollinator-mediated selection may maintain species-specific floral traits.  相似文献   

14.
The role of badges as indicators of contest ability has been previously described. In hummingbirds, the exhibition of a badge is expected to save energy expenditure in agonistic interactions and to favor energy intake. Here, we investigate whether variable supercilium size in the white-eared hummingbird has a role in dominance status signaling. Firstly, 45 hummingbird males were captured and their superciliums were photographed to investigate variation in size and any possible allometric relationships. Secondly, 42 male birds were used to analyze whether the supercilium has a role in dominance status signaling in a dyadic contest. We found that supercilium size varied continuously but that despite variability between individuals, there was no relationship between supercilium size and body size. However, our dyad experiment indicated that birds with larger badges were able to make more visits to the feeders than individuals with smaller badges. We suggest a status signaling function of the supercilium.  相似文献   

15.
Flower Constancy, Insect Psychology, and Plant Evolution   总被引:1,自引:0,他引:1  
 Individuals of some species of pollinating insects tend to restrict their visits to only a few of the available plant species, in the process bypassing valuable food sources. The question of why this flower constancy exists is a rich and important one with implications for the organization of natural communities of plants, floral evolution, and our understanding of the learning processes involved in finding food. Some scientists have assumed that flower constancy is adaptive per se. Others argued that constancy occurs because memory capacity for floral features in insects is limited, but attempts to identify the limitations often remained rather simplistic. We elucidate now different sensory and motor memories from natural foraging tasks are stored and retrieved, using concepts from modern learning science and visual search, and conclude that flower constancy is likely to have multiple causes. Possible constraints favoring constancy are interference sensitivity of short-term memory, and temporal limitations on retrieving information from long-term memory as rapidly as from short-term memory, but further empirical evidence is needed to substantiate these possibilities. In addition, retrieving memories may be slower and more prone to errors when there are several options than when an insect copes with only a single task. In addition to memory limitations, we also point out alternative explanations for flower constancy. We then consider the way in which floral parameters, such as interplant distances, nectar rewards, flower morphology, and floral color (as seen through bees' eyes) affect constancy. Finally, we discuss the implications of pollinator constancy for plant evolution. To date there is no evidence that flowers have diverged to favor constancy, although the appropriate tests may not have yet been conducted. However, there is good evidence against the notion that pollinator constancy is involved in speciation or maintenance of plant species integrity.  相似文献   

16.
Biological invasions can affect the structure and function of ecosystems and threaten native plant species. Since most weeds rely on mutualistic relationships in their new environment, they may act as new competitors for pollinators. Pollinator competition is likely to be density dependent, but it is often difficult to disentangle competition caused by flower quality from effects caused by flower quantity. In order to test the effects of the presence and number of flowers of the invasive weed Bunias orientalis on the insect visitation rates in a native species (Sinapis arvensis), we performed two replacement experiments using plants with standardised flower numbers. The visitation rates in S. arvensis were significantly higher than in B. orientalis and the number of insect visits dropped significantly with increasing density of S. arvensis flowers. These results suggest that intraspecific competition among flowers of S. arvensis is stronger than the competitive effect of alien flowers. As flowers of B. orientalis do not seem to distract visitors from S. arvensis, it is unlikely that pollinator competition between these two plant species plays a crucial role. However, it cannot be excluded that mass blossom stands of B. orientalis may distract flower visitors from native species.  相似文献   

17.
Nectar is secreted in particular rhythms throughout the lifespan of a flower, which allows determining the nectar production dynamics. This paper compares nectar features in Mucuna japira and Mucuna urens describing: dynamics of nectar production, floral response to nectar removal, resorption, nectar sugar composition, and variation in nectar sugar composition. M. japira inflorescence bears 12–21 yellow flowers, which are in anthesis for 7 days, whereas M. urens inflorescence bears 36–54 greenish flowers, but only 1–3 flowers are in anthesis simultaneously that last one night. Nectar volume and sugar concentration were measured, and the amount of sugar was estimated. Qualitative and quantitative nectar sugar composition was determined. Both species had a constant nectar sugar concentration (ca. 10% for M. japira and ca. 16% for M. urens) and secreted high volumes of nectar (ca. 340 μl per flower for M. japira and 310 μl per flower for M. urens), during 5 days for M. japira and 6 h for M. urens, but after the first removal, i.e., when flower opening mechanism is triggered, nectar production stops immediately. Nectar resorption occurred in both species. Nectar sugar composition showed some similarities between the species. Variation in nectar sugar composition occurred in both species. The Mucuna species are dependent on their pollinators to produce fruits and seeds, and they have different strategies to promote the necessary interaction with birds or bats, especially related to nectar and flower characteristics.  相似文献   

18.
To forage effectively amongst flowers, some bee species utilize olfactory cues left by previous visitors in addition to direct assessment of visual cues to identify rewarding flowers. This ability can be more advantageous if the bees can recognize and use scent marks left by heterospecifics, not just marks left by members of their own species. We conducted field experiments to investigate whether the sweat bee Halictus aerarius avoids visiting flowers of trailing water willow Justicia procumbens emptied by other bee species. We found that H. aerarius rejected the flowers visited by both heterospecifics and conspecifics. They also rejected visited flowers artificially replenished with nectar. Our results demonstrate that social bees outside the Apidae can detect marks left on flowers by heterospecifics but that (on this plant species) they are unable to discriminate against flowers by directly detecting nectar volume. H. aerarius exhibited different rejection rates according to the identity of the previous bee species. We suggest that the frequency of rejection responses may depend on the amount of chemical substances left by the previous bee. In general, the use of scent marks left by previous visitors is almost certainly advantageous, enabling foragers to avoid flowers with depleted nectar levels and thereby improving their foraging efficiency.  相似文献   

19.
Nectar guides can enhance pollinator efficiency and plant fitness by allowing pollinators to more rapidly find and remember the location of floral nectar. We tested if a radiating nectar guide around a nectary would enhance the ability of naïve bumble bee foragers to find nectar. Most experiments that test nectar guide efficacy, specifically radiating linear guides, have used guides positioned around the center of a radially symmetric flower, where nectaries are often found. However, the flower center may be intrinsically attractive. We therefore used an off-center guide and nectary and compared “conjunct” feeders with a nectar guide surrounding the nectary to “disjunct” feeders with a nectar guide separated from the nectary. We focused on the innate response of novice bee foragers that had never previously visited such feeders. We hypothesized that a disjunct nectar guide would conflict with the visual information provided by the nectary and negatively affect foraging. Approximately, equal numbers of bumble bees (Bombus impatiens) found nectar on both feeder types. On disjunct feeders, however, unsuccessful foragers spent significantly more time (on average 1.6-fold longer) searching for nectar than any other forager group. Successful foragers on disjunct feeders approached these feeders from random directions unlike successful foragers on conjunct feeders, which preferentially approached the combined nectary and nectar guide. Thus, the nectary and a surrounding nectar guide can be considered a combination of two signals that attract naïve foragers even when not in the floral center.  相似文献   

20.
Fires in the Cerrado savanna are a severe form of disturbance, but some species are capable of resprouting afterwards. It is unknown, however, how and whether post-fire resprouting represents a stressful condition to plants and how their rapid re-growth influences both the production of biochemical compounds, and interactions with mutualistic ants. In this study, we examined the influence of post-fire resprouting on biotic interactions (ant–plant–herbivore relationships) and on plant stress. The study was performed on two groups of the extrafloral nectaried shrub Banisteriopsis campestris (Malpighiaceae); one group was recovering from fire while the other acted as control. With respect to biotic interactions, we examined whether resprouting influenced extrafloral nectar concentration (milligrams per microliter), the abundance of the ant Camponotus crassus and leaf herbivory rates. Plant stress was assessed via fluctuating asymmetry (FA) analysis, which refers to deviations from perfect symmetry in bilaterally symmetrical traits (e.g., leaves) and indicates whether species are under stress. Results revealed that FA, sugar concentration, and ant abundance were 51.7 %, 35.7 % and 21.7 % higher in resprouting plants. Furthermore, C. crassus was significantly associated with low herbivory rates, but only in resprouting plants. This study showed that post-fire resprouting induced high levels of plant stress and influenced extrafloral nectar quality and ant-herbivore relationships in B. campestris. Therefore, despite being a stressful condition to the plant, post-fire resprouting individuals had concentrated extrafloral nectar and sustained more ants, thus strengthening the outcomes of ant–plant mutualism.  相似文献   

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