Are pollinators the agents of selection on flower colour and size in irises?

Plant–pollinator interactions are believed to play a major role in the evolution of floral traits. Flower colour and flower size are important for attracting pollinators, directly influencing reproduction, and thus expected to be under pollinator‐mediated selection. Pollinator‐mediated selection is also proposed to play a role in maintaining flower colour polymorphism within populations. However, pigment concentrations, and thus flower colour, are also under selective pressures independent of pollinators. We quantified phenotypic pollinator‐mediated selection on flower colour and size in two colour polymorphic Iris species. Using female fitness, we estimated phenotypic selection on flower colour and size, and tested for pollinator‐mediated selection by comparing selection gradients between flowers open to natural pollination and supplementary pollinated flowers. In both species, we found evidence for pollen limitation, which set the base for pollinator‐mediated selection. In the colour dimorphic Iris lutescens, while pigment concentration and flower size were found to be under selection, this was independent of pollinators. For the polymorphic Iris pumila, pigment concentration is under selective pressure by pollinators, but only for one colour morph. Our results suggest that pollinators are not the main agents of selection on floral traits in these irises, as opposed to the accepted paradigm on floral evolution. This study provides an opposing example to the largely‐accepted theory that pollinators are the major agent of selection on floral traits.     

Optimal foraging when predation risk increases with patch resources: an analysis of pollinators and ambush predators

Pollinators and their predators share innate and learned preferences for high quality flowers. Consequently, pollinators are more likely to encounter predators when visiting the most rewarding flowers. I present a model of how different pollinator species can maximize lifetime resource gains depending on the density and distribution of predators, as well as their vulnerability to capture by predators. For pollinator species that are difficult for predators to capture, the optimal strategy is to visit the most rewarding flowers as long as predator density is low. At higher predator densities and for pollinators that are more vulnerable to predator capture, the lifetime resource gain from the most rewarding flowers declines and the optimal strategy depends on the predator distribution. In some cases, a wide range of floral rewards provides near‐maximum lifetime resource gains, which may favor generalization if searching for flowers is costly. In other cases, a low flower reward level provides the maximum lifetime resource gain and so pollinators should specialize on less rewarding flowers. Thus, the model suggests that predators can have qualitatively different top‐down effects on plant reproductive success depending on the pollinator species, the density of predators, and the distribution of predators across flower reward levels.     

A meta-analysis of predation risk effects on pollinator behaviour

Flower-visiting animals are constantly under predation risk when foraging and hence might be expected to evolve behavioural adaptations to avoid predators. We reviewed the available published and unpublished data to assess the overall effects of predators on pollinator behaviour and to examine sources of variation in these effects. The results of our meta-analysis showed that predation risk significantly decreased flower visitation rates (by 36%) and time spent on flowers (by 51%) by pollinators. The strength of the predator effects depended neither on predator taxa and foraging mode (sit-and-wait or active hunters) nor on pollinator lifestyle (social vs. solitary). However, predator effects differed among pollinator taxa: predator presence reduced flower visitation rates and time spent on flowers by Squamata, Lepidoptera and Hymenoptera, but not by Diptera. Furthermore, larger pollinators showed weaker responses to predation risk, probably because they are more difficult to capture. Presence of live crab spiders on flowers had weaker effects on pollinator behaviour than presence of dead or artificial crab spiders or other objects (e.g. dead bees, spheres), suggesting that predator crypsis may be effective to some extent. These results add to a growing consensus on the importance of considering both predator and pollinator characteristics from a community perspective.     

Optimality modeling and fitness trade‐offs: when should plants become pollinator specialists?

The assumption that flowers readily evolve specializations for pollination by particular animals has been central to a standard view of pollinator-mediated adaptive divergence in angiosperms. Stebbins' Most Effective Pollinator Principle (MEPP) formalized this assumption in proposing that a plant should always evolve specializations to its most effective pollinator. I argue that the MEPP and its corollaries are unsupported by basic models of phenotypic selection which predict that a plant should evolve greater specialization to a particular pollinator when the marginal fitness gain exceeds the marginal fitness loss from becoming less adapted to all other pollinators. Differences in pollinator effectiveness are neither necessary nor sufficient to predict specialization. Differences in effectiveness certainly can foster floral specialization to the most effective pollinator in some cases, but when adaptation to a relatively ineffective pollinator requires little loss in the fitness contribution of a more effective pollinator, plants may exhibit striking specializations for the less effective pollinator. Recognizing that the effectiveness of pollinators is not tightly coupled to their importance in selecting for phenotypic novelty may resolve the mismatch between floral features that appear to represent clear evolutionary responses to specific pollinators and patterns of flower visitation that often seem generalized. To shed light on agents of selection and the adaptive value of floral traits I argue that we must go beyond measures of pollinator effectiveness to investigate pollinator-mediated fitness trade-offs over a range of floral phenotypes.     

Floral antagonists counteract pollinator‐mediated selection on attractiveness traits in the hummingbird‐pollinated Collaea cipoensis (Fabaceae)

Pollinator‐mediated selection toward larger and abundant flowers is common in naturally pollen‐limited populations. However, floral antagonists may counteract this effect, maintaining smaller‐ and few‐flowered individuals within populations. We quantified pollinator and antagonist visit rates and determined a multiplicative female fitness component from attacked and non‐attacked flowers of the Brazilian hummingbird‐pollinated shrub Collaea cipoensis to determine the selective effects of pollinators and floral antagonists on flower size and number. We predicted that floral antagonists reduce the female fitness component and thus exert negative selective pressures on flower size and number, counteracting the positive effects of pollinators. Pollinators, mainly hummingbirds, comprised 4% of total floral visitation, whereas antagonist ants and bees accounted for 90% of visitation. Nectar‐robbers involved about 99% of floral antagonist visit rates, whereas florivores comprised the remaining 1%. Larger and abundant flowers increased both pollinator and antagonist visit rates and the female fitness component significantly decreased in flowers attacked by nectar‐robbers and florivores in comparison to non‐attacked flowers. We detected that pollinators favored larger‐ and many‐flowered individuals, whereas floral antagonists exerted negative selection on flower size and number. This study confirms that floral antagonists reduce female plant fitness and this pattern directly exerts negative selective pressures on flower size and number, counteracting pollinator‐mediated selection on floral attractiveness traits.     

Do pollinators influence the assembly of flower colours within plant communities?

The co-occurrence of plant species within a community is influenced by local deterministic or neutral processes as well as historical regional processes. Floral trait distributions of co-flowering species that share pollinators may reflect the impact of pollinator preference and constancy on their assembly within local communities. While pollinator sharing may lead to increased visitation rates for species with similar flowers, the receipt of foreign pollen via interspecific pollinator movements can decrease seed set. We investigated the pattern of community flower colour assembly as perceived by native honeybee pollinators within 24 local assemblages of co-flowering Oxalis species within the Greater Cape Floristic Region, South Africa. To explore the influence of pollinators on trait assembly, we assessed the impact of colour similarity on pollinator choices and the cost of heterospecific pollen receipt. We show that flower colour is significantly clustered within Oxalis communities and that this is not due to historical constraint, as flower colour is evolutionarily labile within Oxalis and communities are randomly structured with respect to phylogeny. Pollinator observations reveal that the likelihood of pollinators switching between co-flowering species is low and increases with flower colour similarity. Interspecific hand pollination significantly reduced seed set in the four Oxalis species we investigated, and all were dependant on pollinators for reproduction. Together these results imply that flower colour similarity carries a potential fitness cost. However, pollinators were highly flower constant, and remained so despite the extreme similarity of flower colour as perceived by honeybees. This suggests that other floral traits facilitate discrimination between similarly coloured species, thereby likely resulting in a low incidence of interspecific pollen transfer (IPT). If colour similarity promotes pollinator attraction at the community level, the observed clustering of flower colour within communities might result from indirect facilitative interactions.     

Selection of trait combinations through bee and fly visitation to flowers of Polemonium foliosissimum

Pollinators are known to exert natural selection on floral traits, but the extent to which combinations of floral traits are subject to correlational selection (nonadditive effects of two traits on fitness) is not well understood. Over two years, we used phenotypic manipulations of plant traits to test for effects of flower colour, flower shape and their interaction on rates of pollinator visitation to Polemonium foliosissimum. We also tested for correlational selection based on weighting visitation by the amount of conspecific pollen delivered per visit by each category of insect visitor. Although bumblebees were the presumed pollinators, solitary bees and flies contributed substantially (42%) to pollination. In manipulations of one trait at a time, insects visited flowers presenting the natural colour and shape over flowers manipulated to present artificial mutants with either paler colour or a more open or more tubular flower. When both colour and shape were manipulated in combination, selection on both traits arose, with bumblebees responding mainly to colour and flies responding mainly to shape. Despite selection on both floral traits, in a year with many bumblebees, we saw no evidence for correlational selection of these traits. In a year when flies predominated, fly visitation showed a pattern of correlational selection, but not favouring the natural phenotype, and correlational selection was still not detected for expected pollen receipt. These results show that flower colour and shape are subject to pollinator‐mediated selection and that correlational selection can be generated based on pollinator visitation alone, but provide no evidence for correlational selection specifically for the current phenotype.     

Altitudinal flower size variation correlates with local pollinator size in a bumblebee‐pollinated herb,Prunella vulgaris L. (Lamiaceae)

The influence of locally different species interactions on trait evolution is a focus of recent evolutionary studies. However, few studies have demonstrated that geographically different pollinator‐mediated selection influences geographic variation in floral traits, especially across a narrow geographic range. Here, we hypothesized that floral size variation in the Japanese herb Prunella vulgaris L. (Lamiaceae) is affected by geographically different pollinator sizes reflecting different pollinator assemblages. To evaluate this hypothesis, we posed two questions. (1) Is there a positive correlation between floral length and the proboscis length of pollinators (bumblebees) across altitude in a mountain range? (2) Does the flower–pollinator size match influence female and male plant fitness? We found geographic variation in the assemblage of pollinators of P. vulgaris along an altitudinal gradient, and, as a consequence, the mean pollinator proboscis length also changed altitudinally. The floral corolla length of P. vulgaris also varied along an altitudinal gradient, and this variation strongly correlated with the local pollinator size but did not correlate with altitude itself. Furthermore, we found that the size match between the floral corolla length and bee proboscis length affected female and male plant fitness and the optimal size match (associated with peak fitness) was similar for the female and male fitness. Collectively, these results suggest that pollinator‐mediated selection influences spatial variation in the size of P. vulgaris flowers at a fine spatial scale.     

Predator crypsis enhances behaviourally mediated indirect effects on plants by altering bumblebee foraging preferences

Predators of pollinators can influence pollination services and plant fitness via both consumptive (reducing pollinator density) and non-consumptive (altering pollinator behaviour) effects. However, a better knowledge of the mechanisms underlying behaviourally mediated indirect effects of predators is necessary to properly understand their role in community dynamics. We used the tripartite relationship between bumblebees, predatory crab spiders and flowers to ask whether behaviourally mediated effects are localized to flowers harbouring predators, or whether bees extend their avoidance to entire plant species. In a tightly controlled laboratory environment, bumblebees (Bombus terrestris) were exposed to a random mixture of equally rewarding yellow and white artificial flowers, but foraging on yellow flowers was very risky: bees had a 25 per cent chance of receiving a simulated predation attempt by ‘robotic’ crab spiders. As bees learnt to avoid ‘dangerous’ flowers, their foraging preferences changed and they began to visit fewer yellow flowers than expected by chance. Bees avoided spider-free yellow flowers as well as dangerous yellow flowers when spiders were more difficult to detect (the colour of yellow spiders was indistinguishable from that of yellow flowers). Therefore, this interaction between bee learning and predator crypsis could lead flower species harbouring cryptic predators to suffer from reduced reproductive success.     

The Roles of Colour and Shape in Pollinator Deception in the Orchid Mantis Hymenopus coronatus

The orchid mantis Hymenopus coronatus (Insecta: Mantodea) is a deceptive predator that attracts pollinators as prey. Their resemblance to a flower has given rise to the hypothesis that they are flower mimics. However, floral mimicry as a predatory strategy, and in particular, how predatory floral mimicry functions at a mechanistic level is poorly understood. Two main morphological characteristics are thought to make orchid mantises appear similar to flowers and thus attractive to pollinators: (1) their ‘flower‐like’ white colouration and (2) their ‘petal‐shaped’ expansions of exoskeleton on their mid‐femur and hind femur (femoral lobes). I investigated the contribution of these colour and shape characteristics to pollinator attraction using artificial orchid mantis models. Models with the ‘flower‐like’ white colouration of the orchid mantis had higher rates of pollinator inspection than brown models. Manipulating overall body shape by removing or changing the orientation of the ‘petal‐shaped’ femoral lobes did not affect the attractiveness of models. As certain flower‐like characteristics (symmetry and petals) did not affect the attractiveness of models, pollinators may not necessarily cognitively misclassify orchid mantises as flowers. Rather, mantises may be exploiting sensory biases of their pollinator prey, and their UV‐absorbing white colouration may be sufficient to lure pollinators. The effectiveness of using artificial models established here provides a basis for future research into orchid mantis morphology and the fine‐scale interactions between orchid mantises and pollinators.     

Flower colour adaptation in a mimetic orchid

Although the tremendous variability in floral colour among angiosperms is often attributed to divergent selection by pollinators, it is usually difficult to preclude the possibility that floral colour shifts were driven by non-pollinator processes. Here, we examine the adaptive significance of flower colour in Disa ferruginea, a non-rewarding orchid that is thought to attract its butterfly pollinator by mimicking the flowers of sympatric nectar-producing species. Disa ferruginea has red flowers in the western part of its range and orange flowers in the eastern part--a colour shift that we hypothesized to be the outcome of selection for resemblance to different local nectar-producing plants. Using reciprocal translocations of red and orange phenotypes as well as arrays of artificial flowers, we found that the butterfly Aeropetes tulbaghia, the only pollinator of the orchid, preferred both the red phenotype and red artificial flowers in the west where its main nectar plant also has red flowers, and both the orange phenotype and orange artificial flowers in the east, where its main nectar plant has orange flowers. This phenotype by environment interaction demonstrates that the flower colour shift in D. ferruginea is adaptive and driven by local colour preference in its pollinator.     

Pollinator parasites and the evolution of floral traits

The main selective force driving floral evolution and diversity is plant–pollinator interactions. Pollinators use floral signals and indirect cues to assess flower reward, and the ensuing flower choice has major implications for plant fitness. While many pollinator behaviors have been described, the impact of parasites on pollinator foraging decisions and plant–pollinator interactions have been largely overlooked. Growing evidence of the transmission of parasites through the shared‐use of flowers by pollinators demonstrate the importance of behavioral immunity (altered behaviors that enhance parasite resistance) to pollinator health. During foraging bouts, pollinators can protect themselves against parasites through self‐medication, disease avoidance, and grooming. Recent studies have documented immune behaviors in foraging pollinators, as well as the impacts of such behaviors on flower visitation. Because pollinator parasites can affect flower choice and pollen dispersal, they may ultimately impact flower fitness. Here, we discuss how pollinator immune behaviors and floral traits may affect the presence and transmission of pollinator parasites, as well as how pollinator parasites, through these immune behaviors, can impact plant–pollinator interactions. We further discuss how pollinator immune behaviors can impact plant fitness, and how floral traits may adapt to optimize plant fitness in response to pollinator parasites. We propose future research directions to assess the role of pollinator parasites in plant–pollinator interactions and evolution, and we propose better integration of the role of pollinator parasites into research related to pollinator optimal foraging theory, floral diversity and agricultural practices.     

ECOLOGICAL COSTS OF PLANT RESISTANCE TO HERBIVORES IN THE CURRENCY OF POLLINATION

In this paper, we examine how ecological costs of resistance might be manifested through plant relationships with pollinators. If defensive compounds are incorporated into floral structures or if they are sufficiently costly that fewer rewards are offered to pollinators, pollinators may discriminate against more defended plants. Here we consider whether directional selection for increased resistance to herbivores could be constrained by opposing selection through pollinator discrimination against more defended plants. We used artificial selection to create two populations of Brassica rapa plants that had high and low myrosinase concentrations and, consequently, high and low resistance to flea beetle herbivores. We measured changes in floral characters of plants in both damaged and undamaged states from these populations with different resistances to flea beetle attack. We also measured pollinator visitation to plants, including numbers of pollinators and measures of visit quality (numbers of flowers visited and time spent per flower). Damage from herbivores resulted in reduced petal size, as did selection for high resistance to herbivores later in the plant lifetime. In addition, floral display (number of open flowers) was also altered by an interaction between these two effects. Changes in floral traits translated into overall greater use of low-resistance, undamaged plants based on total amount of time pollinators spent foraging on plants. Total numbers of pollinators attracted to plants did not differ among treatments; however, pollinators spent significantly more time per flower on plants from the low-resistance population and tended to visit more flowers on these plants as well. Previous work by other investigators on the same pollinator taxa has shown that longer visit times are associated with greater male and female plant fitness. Because initial numbers of pollinators did not differ between selection regimes, palatability and/or amount of rewards offered by high- and low-resistance populations are likely to be responsible for these patterns. During periods of pollinator limitation, less defended plants may have a selective advantage and pollinator preferences may mediate directional selection imposed by herbivores. In addition, if pollinator preferences limit seed set in highly defended plants, then lower seed set previously attributed to allocation costs of defense may also reflect greater pollinator limitation in these plants relative to less defended plants.     

Effects of experimental manipulation of inflorescence size on pollination and pre-dispersal seed predation in the hummingbird-pollinated plant Ipomopsis aggregata

Large floral displays should theoretically provide advantages to plants through increased pollinator visitation and resulting fruit and seed set. However empirical tests of the response of pollinators to floral display size have been limited by a lack of direct experimentation, and the results of such studies have been equivocal. In addition, other selective agents such as pre-dispersal seed predators might modulate effects of floral display on pollination. By artificially altering flower number, we examined the direct effects of floral display in the monocarpic herb, Ipomopsis aggregata (Polemoniaceae), on visitation rates by broad-tailed and rufous hummingbird pollinators, as well destruction of fruits by a pre-dispersal seed predator (Hylemya: Anthomyiidae). In addition, we quantified the ultimate effects of flower number on female reproductive success. Plants with larger floral displays were most likely to be visited first in any given foraging bout (P < 0.01). As expected, plants with more flowers received more total flower visits. However, we found no gain in the proportion of flowers visited for many- versus few-flowered plants, or the total number of approaches/hour. In fact, a significantly greater percentage of flowers were visited on few-flowered plants. Plants did not compensate for our reduction in flowers by increasing investment in the number or proportion of flowers that set fruit, the number of seeds/fruit, or seed weight. Pre-dispersal seed predation was greater for many- than for few-flowered plants (P < 0.001), but this did not offset the potential fitness gains of producing large displays. Our data support the hypothesis that large floral displays function primarily in long-distance attraction of pollinators, and enhance maternal success. Received: 21 March 1996 / Accepted: 24 October 1996     

Pollinator response to flower color polymorphism and floral display in a plant with a single-locus floral color polymorphism: consequences for plant reproduction

Variation in flower color, particularly polymorphism, in which two or more different flower color phenotypes occur in the same population or species, may be affected or maintained by mechanisms that depend on pollinators. Furthermore, variation in floral display may affect pollinator response and plant reproductive success through changes in pollinator visitation and availability of compatible pollen. To asses if flower color polymorphism and floral display influences pollinator preferences and movements within and among plants and fitness-related variables we used the self-incompatible species Cosmos bipinnatus Cav. (Asteraceae), a model system with single-locus flower color polymorphism that comprises three morphs: white (recessive homozygous), pink (heterozygous co-dominate), and purple (dominant homozygous) flowers. We measured the preferences of pollinators for each morph and constancy index for each pollinator species, pollination visitation rate, floral traits, and female fitness measures. Flower color morphs differed in floral trait measures and seed production. Pollinators foraged nonrandomly with respect to flower color. The most frequent morph, the pink morph, was the most visited and pollinators exhibited the highest constancy for this morph. Moreover, this morph exhibited the highest female fitness. Pollinators responded strongly to floral display size, while probed more capitulums from plants with large total display sizes, they left a great proportion of them unvisited. Furthermore, total pollinator visitation showed a positive relation with female fitness. Results suggest that although pollinators preferred the heterozygous morph, they alternate indiscriminately among morphs making this polymorphism stable.     

Pollinator‐mediated phenotypic selection does not always modulate flower size and number in the large‐flowered Mediterranean shrub Cistus ladanifer (Cistaceae)

Flower size and number usually evolve under pollinator‐mediated selection. However, hot, dry environments can also modulate display, counteracting pollinator attraction. Increased pollen deposition on larger flower displays may not involve higher female fitness. Consequently, stressful conditions may constrain flower size, favouring smaller‐sized flowers. The large‐flowered, self‐incompatible Mediterranean shrub Cistus ladanifer was used to test that: (1) this species suffers pollen limitation; (2) pollinators are spatially–temporally variable and differentially visit plants with more/larger flowers; (3) increased visits enhance reproduction under pollen limitation; (4) stressful conditions reduce female fitness of larger displays; and (5) phenotypic selection on floral display is not just pollinator‐mediated. We evaluated pollen limitation, related floral display to pollinator visits and fruit and seed production and estimated phenotypic selection. Flower size was 7.2–10.5 cm and varied spatially–temporally. Visitation rates (total visits/50 min) ranged from 0.26 to 0.43 and increased with flower size. Fruit set averaged 80% and seed number averaged 855, but only fruit set varied between populations and years. Selection towards larger flowers was detected under conditions of pollen limitation. Otherwise, we detected stabilizing selection on flower size and negative selection on flower number. Our results suggest that selection on floral display is not only pollinator‐dependent through female fitness in C. ladanifer. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 540–555.     

No evidence for a role of pollinator discrimination in causing selection on flower size through female reproduction

The preference for certain floral phenotypes by flower visiting animals may fuel the evolution of floral traits because variation in flower visitation rates may lead to fitness variation within a population. Here, I examine the importance of flower size for pollinator visitation rate, seed set, and seed mass in two alpine populations of the insect-pollinated herb Ranunculus acris L. during two seasons. There was no pollen limitation of seed set or mass. Pollinators discriminated strongly against flowers experimentally reduced in size. Despite this, there were no signs of any significant impact of flower size on female reproductive success. The results show that although pollinators discriminate strongly among floral phenotypes, this may not always result in female fitness differences within a population because seed set or mass is not limited by pollen availability alone. Probably abiotic environmental constraints prevent plants with high pollinator visitation from capitalizing on the high pollen deposition.     

Evolution of honest reward signal in flowers

Some flowering plants signal the abundance of their rewards by changing their flower colour, scent or other floral traits as rewards are depleted. These floral trait changes can be regarded as honest signals of reward states for pollinators. Previous studies have hypothesized that these signals are used to maintain plant-level attractiveness to pollinators, but the evolutionary conditions leading to the development of honest signals have not been well investigated from a theoretical basis. We examined conditions leading to the evolution of honest reward signals in flowers by applying a theoretical model that included pollinator response and signal accuracy. We assumed that pollinators learn floral traits and plant locations in association with reward states and use this information to decide which flowers to visit. While manipulating the level of associative learning, we investigated optimal flower longevity, the proportion of reward and rewardless flowers, and honest- and dishonest-signalling strategies. We found that honest signals are evolutionarily stable only when flowers are visited by pollinators with both high and low learning abilities. These findings imply that behavioural variation in learning within a pollinator community can lead to the evolution of an honest signal even when there is no contribution of rewardless flowers to pollinator attractiveness.     

Floral organs act as environmental filters and interact with pollinators to structure the yellow monkeyflower (Mimulus guttatus) floral microbiome

Assembly of microbial communities is the result of neutral and selective processes. However, the relative importance of these processes is still debated. Microbial communities of flowers, in particular, have gained recent attention because of their potential impact to plant fitness and plant‐pollinator interactions. However, the role of selection and dispersal in the assembly of these communities remains poorly understood. Here, we evaluated the role of pollinator‐mediated dispersal on the contribution of neutral and selective processes in the assembly of floral microbiomes of the yellow monkeyflower (Mimulus guttatus). We sampled floral organs from flowers in the presence and absence of pollinators within five different serpentine seeps in CA and obtained 16S amplicon data on the epiphytic bacterial communities. Consistent with strong microenvironment selection within flowers we observed significant differences in community composition across floral organs and only a small effect of geographic distance. Pollinator exposure affected the contribution of environmental selection and depended on the rate and intimacy of interactions with flower visitors. This study provides evidence of the importance of dispersal and within‐flower heterogeneity in shaping epiphytic bacterial communities of flowers, and highlights the complex interplay between pollinator behaviour, environmental selection and additional abiotic factors in shaping the epiphytic bacterial communities of flowers.     

毛茛状金莲花不同花期的花特征和访花昆虫的变化及表型选择

为了研究植物生长季内开花时间对花特征表型选择的影响,我们以青藏高原东缘高寒草地的毛茛状金莲花Trollius ranunculoides)为实验材料,在生长季内不同开花时间(花前期、花末期)测定花特征,观察访花昆虫的类群和访花频率,生长季结束后收集种子.根据昆虫访花的喜好和季节内类群与访花频率的变化,分析了不同开花时间毛茛状金莲花的花特征与昆虫的选择;并用种子产量表示雌性适合度,估计了毛茛状金莲花的花特征在不同开花时间所受的表型选择.结果表明:不同花期植物的花特征有显著差异,相应的访花昆虫的类群和频率也存在差异,不同类群昆虫访花喜好也不一样.蜂喜好花瓣和花萼较宽、花茎短和花茎数少的个体,这正符合花前期的特征,因而蜂的访花频率在花前期较高;蝇对花特征没有明显的偏好.而通过雌性适合度估计毛茛状金莲花花特征所受的表型选择则是:花前期,花茎较长和花茎数多的植株适合度大;花末期,花茎数多的植株适合度大.我们的研究表明:在植物生长季,花期的分化伴随着传粉昆虫活动的变化.不同花期,访花昆虫的变化可能对植物花特征的分化起了至关重要的作用.但是访花昆虫对花特征的选择与通过雌性适合度估计植物受到的选择不尽相同,这可能是由于其他因素造成的.