Quantifying hummingbird preference for floral trait combinations: The role of selection on trait interactions in the evolution of pollination syndromes

Darwin recognized the flower's importance for the study of adaptation and emphasized that the flower's functionality reflects the coordinated action of multiple traits. Here we use a multitrait manipulative approach to quantify the potential role of selection acting on floral trait combinations underlying the divergence and maintenance of three related North American species of Silene (Caryophyllaceae). We artificially generated 48 plant phenotypes corresponding to all combinations of key attractive traits differing among the three Silene species (color, height, inflorescence architecture, flower orientation, and corolla‐tube width). We quantified main and interaction effects of trait manipulation on hummingbird visitation preference using experimental arrays. The main effects of floral display height and floral orientation strongly influenced hummingbird visitation, with hummingbirds preferring flowers held high above the ground and vertically to the sky. Hummingbirds also prefer traits in a nonadditive manner as multiple two‐way and higher order interaction effects were important predictors of hummingbird visitation. Contemporary trait combinations found in hummingbird pollinated S. virginica are mostly preferred. Our study demonstrates the likelihood of pollination syndromes evolving due to selection on trait combinations and highlights the importance of trait interactions in understanding the evolution of complex adaptations.     

Floral trait evolution associated with shifts between insect and wind pollination in the dioecious genus Leucadendron (Proteaceae)

Transitions between animal and wind pollination have occurred in many lineages and have been linked to various floral modifications, but these have seldom been assessed in a phylogenetic framework. In the dioecious genus Leucadendron (Proteaceae), transitions from insect to wind pollination have occurred at least four times. Using analyses that controlled for relatedness among Leucadendron species, we investigated how these transitions shaped the evolution of floral structural and signaling traits, including the degree of sexual dimorphism in these traits. Pollen grains of wind‐pollinated species were found to be smaller, more numerous, and dispersed more efficiently in wind than were those of insect‐pollinated species. Wind‐pollinated species also exhibited a reduction in spectral contrast between showy subtending leaves and background foliage, reduced volatile emissions, and a greater degree of sexual dimorphism in color and scent. Uniovulate flowers and inflorescence condensation are conserved ancestral features in Leucadendron and likely served as exaptations in shifts to wind pollination. These results offer insights into the key modifications of male and female floral traits involved in transitions between insect and wind pollination.     

Predicting the pathway to wind pollination: heritabilities and genetic correlations of inflorescence traits associated with wind pollination in Schiedea salicaria (Caryophyllaceae)

The transition from biotic to abiotic pollination was investigated using Schiedea, a genus exhibiting a remarkable diversity of inflorescence architecture associated with pollination biology. Heritabilities and genetic correlations of inflorescence traits were estimated in gynodioecious Schiedea salicaria (Caryophyllaceae), a species that has likely undergone a recent transition to wind-pollination. Using a partial diallel crossing design, significant narrow-sense heritabilities were detected for inflorescence condensation (h2 = 0.56 to 0.68 in the two sexes) and other traits related to the extent of wind pollination in Schiedea species. Heritabilities were generally higher in hermaphrodites than in females. Strong genetic correlations may constrain the evolution of some inflorescence traits that facilitate wind pollination, such as simultaneous shortening of inflorescence length and elongation of the subtending internode. The presence of significant narrow-sense heritabilities for traits associated with wind pollination suggests, however, that selection for more effective wind pollination in the windy, pollinator-limited environments where S. salicaria grows could lead to the evolution of the highly condensed inflorescences characteristic of other wind-pollinated species of Schiedea.     

Neither insects nor wind: ambophily in dioecious Chamaedorea palms (Arecaceae)

Pollination of Neotropical dioecious trees is commonly related to generalist insects. Similar data for non‐tree species with separated genders are inconclusive. Recent studies on pollination of dioecious Chamaedorea palms (Arecaceae) suggest that species are either insect‐ or wind‐pollinated. However, the wide variety of inflorescence and floral attributes within the genus suggests mixed pollination mode involving entomophily and anemophily. To evaluate this hypothesis, we studied the pollination of Chamaedorea costaricana, C. macrospadix, C. pinnatifrons and C. tepejilote in two montane forests in Costa Rica. A complementary morphological analysis of floral traits was carried out to distinguish species groups within the genus according to their most probable pollination mechanism. We conducted pollinator exclusion experiments, field observations on visitors to pistillate and staminate inflorescences, and trapped airborne pollen. A cluster analysis using 18 floral traits selected for their association with wind and insect pollination syndromes was carried out using 52 Chamaedorea species. Exclusion experiments showed that both wind and insects, mostly thrips (Thysanoptera), pollinated the studied species. Thrips used staminate inflorescences as brood sites and pollinated pistillate flowers by deception. Insects caught on pistillate inflorescences transported pollen, while traps proved that pollen is wind‐borne. Our empirical findings clearly suggest that pollination of dioecious Chamaedorea palms is likely to involve both insects and wind. A cluster analysis showed that the majority of studied species have a combination of floral traits that allow for both pollination modes. Our pollination experiments and morphological analysis both suggest that while some species may be completely entomophilous or anemophilous, ambophily might be a common condition within Chamaedorea. Our results propose a higher diversity of pollination mechanisms of Neotropical dioecious species than previously suggested.     

Covariance and decoupling of floral and vegetative traits in nine Neotropical plants: a re-evaluation of Berg's correlation-pleiades concept

Nearly forty years ago R. L. Berg proposed that plants with specialized pollination ecology evolve genetic and developmental systems that decouple floral morphology from phenotypic variation in vegetative traits. These species evolve separate floral and vegetative trait clusters, or as she termed them, "correlation pleiades." The predictions of this hypothesis have been generally supported, but only a small sample of temperate-zone herb and grass species has been tested. To further evaluate this hypothesis, especially its applicability to plants of other growth forms, we examined the patterns of phenotypic variation and covariation of floral and vegetative traits in nine species of Neotropical plants. We recognized seven specific predictions of Berg's hypothesis. Our results supported some predictions but not others. Species with specialized pollination systems usually had floral traits decoupled (weak correlation; Canna and Eichornia) or buffered (relationship with shallow proportional slope; Calathea and Canna) from variation in vegetative traits. However, the same trend was also observed in three species with unspecialized pollination systems (Echinodorus, Muntingia, and Wedelia). One species with unspecialized pollination (Croton) and one wind-pollinated species (Cyperus) showed no decoupling or buffering, as predicted. While species with specialized pollination usually showed lower coefficients of variation for floral traits than vegetative traits (as predicted), the same was also true of species with unspecialized or wind pollination (unlike our prediction). Species with specialized pollination showed less variation in floral traits than did species with unspecialized or wind pollination, as predicted. However, the same was true of the corresponding vegetative traits, which was unexpected. Also in contrast to our prediction, plants with specialized pollination systems did not exhibit tighter phenotypic integration of floral characters than did species with generalized pollination systems. We conclude that the patterns of morphological integration among floral traits and between floral and vegetative traits tend to be species specific, not easily predicted from pollination ecology, and generally more complicated than R. L. Berg envisaged.     

Relationships between floral characters, pollination mechanisms, life forms, and habitats in Araceae

The floral traits of the inflorescences of angiosperms have coevolved to ensure and maximize pollination success. Other factors believed to influence floral architecture are external (for example, ecological) to the inflorescence. In order to understand the relationships between such factors and floral characters, 12 floral traits were measured in 54 species of Araceae. An analysis was performed to determine how these traits are linked to the following: (1) self-pollination capacity; (2) life form (evergreen versus seasonally dormant); (3) climatic conditions; and (4) type of pollinator (i.e. flies, bees, or beetles). A significant difference was found between the pollen to ovule ratio of the species able to self-pollinate and those unable to self-pollinate. Evergreen and tropical aroids produced a larger number of gametes than did seasonally dormant and temperate taxa. Finally, several floral traits, such as pollen volume and number, number of female flowers, and flower sexual type (unisexual or bisexual), showed clear differences between the three pollinator types. Variations in floral traits between the different life forms and climatic conditions are discussed with respect to pollination efficiency and properties of the growing season. The pollen to ovule ratio cannot be considered as an accurate indicator of breeding systems in aroids because of the particular pollination ecology of the family.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 29–42.     

Wind pollination, sexual dimorphism, and changes in floral traits of Schiedea (Caryophyllaceae)

Both changes in sex allocation and pollination mode may promote the separation of sexes in plant populations. Simultaneous evolution of wind pollination and dimorphism has occurred in Schiedea, where species with different female frequencies provide an opportunity to observe the effect of wind pollination on sex allocation and floral morphology. Differences among species in the ratio of anther to ovary volume were not the result of sex allocation trade-offs, but instead resulted from production of vestigial stamens in females; there were no changes in ovary volume in males and hermaphrodites (MH) of dimorphic species. Relative to hermaphroditic species, dimorphic species had more condensed inflorescences, a pattern often associated with wind pollination. Within dimorphic species, MH had longer filament lengths than females, and females had longer stigmas than MHs. These traits are characteristic of wind pollination, but there was no relationship between the degree of sexual dimorphism and female frequency. Ovary volume and ovule number and size had positive phenotypic correlations between females and MH of dimorphic species, making sex specialization more difficult. In dimorphic Schiedea species, selection for wind pollination may have a greater effect on floral traits than trade-offs in allocation between male and female function.     

Variation in floral morphology within a population of Aster hispidus var. tubulosus (Asteraceae,Astereae)

The family Asteraceae has a particular inflorescence, the capitulum, consisting of ray florets and disc florets. The ray florets function as petals that attract pollinators. Marked variation in the ray floret morphology is known in a natural population of Aster hispidus var. tubulosus (Asteraceae). We analyzed the variation and found two distinct types in the ray florets, the long tubular ray floret and the ligulate ray floret. In this species, therefore, the variation in floral morphology among capitula, each of which is the basic pollination unit, is caused by the variation in the composition of the two ray floret types among capitula. We evaluated the sources of the observed variation in the floral morphology among capitula within a population using a hierarchical analysis that separated within‐individual (i.e. among capitula within each individual) and between‐individual components of the variation. We found that the main source of the variation lay at the between‐individual level, not at the between‐capitulum level nested within individuals. This finding will provide the basic knowledge that enables future study exploring whether the between‐individual variation in floral morphology caused by the compositional variation of the ray floret types leads to differential pollination success of individual plants in species of Asteraceae.     

Transition from wind pollination to insect pollination in sedges: experimental evidence and functional traits

Transitions from wind pollination to insect pollination were pivotal to the radiation of land plants, yet only a handful are known and the trait shifts required are poorly understood. We tested the hypothesis that a transition to insect pollination took place in the ancestrally wind-pollinated sedges (Cyperaceae) and that floral traits modified during this transition have functional significance. We paired putatively insect-pollinated Cyperus obtusiflorus and Cyperus sphaerocephalus with related, co-flowering, co-occurring wind-pollinated species, and compared pairs in terms of pollination mode and functional roles of floral traits. Experimentally excluding insects reduced seed set by 56-89% in putatively insect-pollinated species but not in intermingled wind-pollinated species. The pollen of putatively insect-pollinated species was less motile in a wind tunnel than that of wind-pollinated species. Bees, beetles and flies preferred inflorescences, and color-matched white or yellow models, of putatively insect-pollinated species over inflorescences, or color-matched brown models, of wind-pollinated species. Floral scents of putatively insect-pollinated species were chemically consistent with those of other insect-pollinated plants, and attracted pollinators; wind-pollinated species were unscented. These results show that a transition from wind pollination to insect pollination occurred in sedges and shed new light on the function of traits involved in this important transition.     

Identification of major quantitative trait loci underlying floral pollination syndrome divergence in Penstemon

Distinct floral pollination syndromes have emerged multiple times during the diversification of flowering plants. For example, in western North America, a hummingbird pollination syndrome has evolved more than 100 times, generally from within insect-pollinated lineages. The hummingbird syndrome is characterized by a suite of floral traits that attracts and facilitates pollen movement by hummingbirds, while at the same time discourages bee visitation. These floral traits generally include large nectar volume, red flower colour, elongated and narrow corolla tubes and reproductive organs that are exerted from the corolla. A handful of studies have examined the genetic architecture of hummingbird pollination syndrome evolution. These studies find that mutations of relatively large effect often explain increased nectar volume and transition to red flower colour. In addition, they suggest that adaptive suites of floral traits may often exhibit a high degree of genetic linkage, which could facilitate their fixation during pollination syndrome evolution. Here, we explore these emerging generalities by investigating the genetic basis of floral pollination syndrome divergence between two related Penstemon species with different pollination syndromes—bee-pollinated P. neomexicanus and closely related hummingbird-pollinated P. barbatus. In an F₂ mapping population derived from a cross between these two species, we characterized the effect size of genetic loci underlying floral trait divergence associated with the transition to bird pollination, as well as correlation structure of floral trait variation. We find the effect sizes of quantitative trait loci for adaptive floral traits are in line with patterns observed in previous studies, and find strong evidence that suites of floral traits are genetically linked. This linkage may be due to genetic proximity or pleiotropic effects of single causative loci. Interestingly, our data suggest that the evolution of floral traits critical for hummingbird pollination was not constrained by negative pleiotropy at loci that show co-localization for multiple traits.     

Flowering dynamics in Arum italicum (Araceae): relative role of inflorescence traits, flowering synchrony, and pollination context on fruit initiation

We studied the relative role of inflorescence traits, flowering synchrony, and pollination context for infructescence and fruit initiation in two Spanish populations of Arum italicum, a species in which inflorescences are the pollination unit. In this species, a specialized inflorescence organ, the appendix, is important for pollinator attraction. However, the short floral longevity and the production of mostly one inflorescence per plant make its pollination potentially dependent on strong flowering synchrony and on external factors not controlled by the plant (the pollination context). The flowering period in both sites lasted >3 mo. Day-to-day variation in simultaneous antheses was high, and 11-50% of antheses occurred on days during which no pollen donor was present. Inflorescence traits, flowering synchrony, and between-plant distance all influenced infructescence and fruit initiation, but their relative importance differed between sites. In one large population, infructescence initiation was positively related to inflorescence traits; in a smaller population infructescence initiation increased with the number of donor inflorescences. In both sites, percentage of fruits initiated per infructescence was dependent on a combination of inflorescence traits, flowering synchrony, and between-plant distance. Plants producing 2-4 inflorescences had higher probability of infructescence initiation and overlapped their antheses with more plants than single-inflorescence ones.     

北热带喀斯特森林木本植物花性状及其生境分异

植物花性状的多样化是植物长期进化及自然选择的结果, 不同植物种间花性状的变异与生境存在一定的相关性。北热带喀斯特季节性雨林具有生境异质性强、群落结构复杂、特有成分丰富等特点, 分析该森林植物性状的变化特征及其与生境的关联性, 有助于理解物种共存、协同进化过程和对生境的适应, 可为阐明喀斯特植物的生态适应性、理解生物多样性维持机理提供依据。本研究在弄岗15 ha森林动态监测样地木本植物开花相对集中的时间段进行, 记录并分析了21个物种花性状的变化。根据物种空间分布及其与生境的关联特性, 将21个物种分成偏好谷底、山坡、山顶的3种类型, 分析了这3种类型植物的花性状差异; 另外又根据花性状对21个物种进行聚类分析, 探讨了聚类分组结果和根据偏好生境分组结果的异同。结果表明: 花大小、花色鲜艳度均与物种优势度存在显著的负相关关系, 花小而不鲜艳的物种在群落中更有优势, 表现出更好的适应性; 不同偏好生境的植物花色明度有显著差异, 其他的花性状差异不显著; 聚类分组与根据偏好生境的分组有较好的一致性, 反映出物种的花性状变化响应了生境的变化。综上结果, 我们认为北热带喀斯特季节性雨林植物花性状与物种的优势度及生境条件有密切联系, 在该区域中生境对花性状的影响可能比传粉者更为深刻。     

Intensified wind pollination mediated by pollen dimorphism after range expansion in an ambophilous biennial Aconitum gymnandrum

Pollination systems and associated floral traits generally differ between core and marginal populations of a species. However, such differences are rarely examined in plants with a mixed wind‐ and bumblebee‐pollination system, and the role of wind pollination during range expansion in ambophilous plants remains unclear. We compared floral traits and the contributions of bumblebee and wind pollination in refugium and marginal populations of the ambophilous plant Aconitum gymnandrum. We found that most floral traits differed between the two populations, and those traits associated with the shift to wind pollination were pronounced in the marginal population. Bumblebee visitation rates varied significantly, but were generally low in the marginal population. Wind pollination occurred in both populations, and the efficiency was lower than that of bumblebee pollination. Two types of pollen grains, namely round and fusiform pollen, were transported to a stigma by bumblebees and wind, but fusiform pollen contributed to wind pollination to a larger degree, especially in the marginal population. Our results suggest that wind pollination was enhanced by pollen dimorphism in the marginal population of A. gymnandrum, and wind pollination may provide reproductive assurance when bumblebee activity is unpredictable during range expansion, indicating that ambophily is stable in this species and shift in pollination system could be common when plants colonize new habitats.     

Branch length mediates flower production and inflorescence architecture of Fouquieria splendens (ocotillo)

The capacity of individual branches to store water and fix carbon can have profound effects on inflorescence size and architecture, thus on floral display, pollination, and fecundity. Mixed regression was used to investigate the relation between branch length, a proxy for plant resources, and floral display of Fouquieria splendens (ocotillo), a woody, candelabraform shrub of wide distribution in arid North America. Long branches produced three times as many flowers as short branches, regardless of overall plant size. Long branches also had more complex panicles with more cymes and cyme types than short branches; thus, branch length also influenced inflorescence architecture. Within panicles, increasing the number of cymes by one unit added about two flowers, whereas increasing the number of cyme types by one unit added about 21 flowers. Because flower production is mediated by branch length, and because most plants have branches of various lengths, the floral display of individual plants necessarily encompasses a wide range of inflorescence size and structure.     

In nomen omen: the effect of flower distance on female reproductive success of the lax-flowered orchid Anacamptis laxiflora (Orchidaceae)

论花距对花序松散的倒距兰(Anacamptis laxiflora)雌花繁殖成功率的影响 对多花物种而言，花序结构是塑造植物吸引力的首要因素。本研究旨在揭示花序性状对植物雌花繁殖成功率的影响，以及倒距兰(Anacamptis laxiflora，一种采用欺骗性授粉策略的陆生物种)花序上的授粉规律。我们还探究了花序性状与雌花繁殖成功率之间的关系是否会因为周围植被的高度和/或种群密度的影响而改变。我们在一个天然的倒距兰种群中划定了实验区块，对每个区块内的植株个体进行计数，并将各区块按低密度和高密度进行分类；随后，在部分区块内，我们人工清除了植株周围的杂草，从而产生了同等数量的高草丛区块和低草丛区块。我们统计了这些区块中的花序性状和雌花繁殖成功率(即果实数量及其在花序上的位置)，使用广义线性混合模型(GLMMs)对统计数据进行了分析，并计算出了选择梯度。我们发现所研究的所有花序性状均对雌花繁殖成功率造成了影响。特别是GLMMs的结果表明，“平均花距”是对繁殖成功规律的最佳预测因子。我们所研究的花序性状受到显著的正向选择作用，但是这些选择趋势与周围植被的高度和种群密度都紧密相关，表明当地环境背景对选择规律的形成有显著影响。雌花繁殖成功率与花朵在花序上的位置无关，这说明传粉者在花序上随机选择花朵，未见其对花序特定部位的偏好。本项研究突显出花序性状对于多花、具有欺骗性的兰花雌花繁殖成功的重要性，并证实了环境背景在调节传粉者介导的选择规律中的主导作用。     

Multiple aspects of the selfing syndrome of the morning glory Ipomoea lacunosa evolved in response to selection: A Qst‐Fst comparison

The frequent transition from outcrossing to selfing in flowering plants is often accompanied by changes in multiple aspects of floral morphology, termed the “selfing syndrome.” While the repeated evolution of these changes suggests a role for natural selection, genetic drift may also be responsible. To determine whether selection or drift shaped different aspects of the pollination syndrome and mating system in the highly selfing morning glory Ipomoea lacunosa, we performed multivariate and univariate Qst‐Fst comparisons using a wide sample of populations of I. lacunosa and its mixed‐mating sister species Ipomoea cordatotriloba. The two species differ in early growth, floral display, inflorescence traits, corolla size, nectar, and pollen number. Our analyses support a role for natural selection driving trait divergence, specifically in corolla size and nectar traits, but not in early growth, display size, inflorescence length, or pollen traits. We also find evidence of selection for reduced herkogamy in I. lacunosa, consistent with selection driving both the transition in mating system and the correlated floral changes. Our research demonstrates that while some aspects of the selfing syndrome evolved in response to selection, others likely evolved due to drift or correlated selection, and the balance between these forces may vary across selfing species.     

Wind of change: new insights on the ecology and evolution of pollination and mating in wind-pollinated plants

Background

The rich literature that characterizes the field of pollination biology has focused largely on animal-pollinated plants. At least 10 % of angiosperms are wind pollinated, and this mode of pollination has evolved on multiple occasions among unrelated lineages, and hence this discrepancy in research interest is surprising. Here, the evolution and functional ecology of pollination and mating in wind-pollinated plants are discussed, a theoretical framework for modelling the selection of wind pollination is outlined, and pollen capture and the occurrence of pollen limitation in diverse wind-pollinated herbs are investigated experimentally.

Scope and Conclusions

Wind pollination may commonly evolve to provide reproductive assurance when pollinators are scarce. Evidence is presented that pollen limitation in wind-pollinated plants may not be as common as it is in animal-pollinated species. The studies of pollen capture in wind-pollinated herbs demonstrate that pollen transfer efficiency is not substantially lower than in animal-pollinated plants as is often assumed. These findings challenge the explanation that the evolution of few ovules in wind-pollinated flowers is associated with low pollen loads. Floral and inflorescence architecture is crucial to pollination and mating because of the aerodynamics of wind pollination. Evidence is provided for the importance of plant height, floral position, and stamen and stigma characteristics in promoting effective pollen dispersal and capture. Finally, it is proposed that geitonogamous selfing may alleviate pollen limitation in many wind-pollinated plants with unisexual flowers.Key words: Wind pollination, reproductive assurance, pollen limitation, geitonogamy, sex allocation, inflorescence architecture, mating systems     

WHY DOES THE SIZE OF REPRODUCTIVE STRUCTURES DECLINE THROUGH TIME IN HYDROPHYLLUM APPENDICULATUM (HYDROPHYLLACEAE)?: DEVELOPMENTAL CONSTRAINTS VS. RESOURCE LIMITATION

A commonly observed pattern in plants is that the size of some reproductive structures declines through a single flowering season. The experimental design of this study allowed me to determine whether architectural constraints or resource limitation is responsible for the seasonal declines in flower size, inflorescence size, and seed weight in the biennial Hydrophyllum appendiculatum (Hydrophyllaceae). Plant resource status was manipulated by varying the number of inflorescences pollinated (two vs. all). Architectural constraints were inferred by comparing the rates of decline in size of reproductive structures in plants that underwent either high or low pollination intensity. The data demonstrated that the relative importance of resource limitation and developmental constraints is trait-specific. Even when not resource stressed, all traits exhibited a seasonal decline, demonstrating that ontogenetic changes occurred in plant architecture. Yet, for two traits, inflorescence size and seed weight, the temporal decline was significantly more severe for heavily pollinated plants. In contrast, the decline in flower size through time was identical in both pollination treatments. Thus, the size of flowers is buffered from the resource status of the plant. These findings illustrate the importance of considering the physical limitations of plant architecture when investigating patterns of plant reproduction.     

INFLORESCENCE ARCHITECTURE: HOW FLOWER NUMBER,ARRANGEMENT, AND PHENOLOGY AFFECT POLLINATION AND FRUIT-SET

The adaptive significance of different types of inflorescences in flowering plants has been largely ignored. The few published studies investigating adaptive aspects of floral displays suggest that numbers of flowers and their arrangement in space and time determine levels of pollination and fruit-set in natural populations. The frequently conflicting demands placed on inflorescence architecture have led to an evolutionary compromise that maximizes the genetic contribution of an individual plant to the next generation. These conflicting demands include pollinator attraction vs. ovary competition, fruit dispersal vs. fruit predation, and reproductive vs. vegetative resource allocation. In most cases, the inflorescence size most successful in fruit production is also the most frequent in natural populations. In addition to quantity of offspring, inflorescence architecture affects, and in turn is affected by, the quality of offspring that result from selfing vs. outcrossing.     

蝶形花亚科植物花部适应机制与传粉系统

蝶形花亚科(Papilionoideae)植物种类丰富、繁育系统多样,与其传粉者关系密切,主要表现在花部适应与传粉系统的形成。蝶形花形态复杂,为完全花,花萼、花瓣5基数,花萼呈箭头状,相邻花瓣螺旋轮生,旗瓣较大位于最外边,两片翼瓣紧贴旗瓣着生并包裹两片合生的龙骨瓣,龙骨瓣内包裹着雌、雄蕊,雌蕊位于正中央,雄蕊轮生,构成二体雄蕊(多数9+1,少数5+5)。对蝶形花亚科植物的花部特征、传粉功能群、酬物与传粉系统构建进行了回顾,重点论述了蝶形花形态和化学组成与传粉系统的进化关系以及花粉呈现机制。其中,泛化的传粉系统以蜂媒传粉为主,同时存在鸟媒传粉、蝙蝠传粉和松鼠传粉等方式,花部结构和传粉者的相互选择和相互适应推动了传粉系统的演化。花粉释放是体现蝶形花植物与其传粉功能群相互作用的重要方面,二者协同进化形成了以下4种花粉呈现机制:弹花机制、活塞机制、瓣膜机制和毛刷机制。蝶形花的花部特征与传粉功能群的相互作用形成相应的传粉系统,而传粉系统是执行传粉功能的重要集合体,各个构件的相互协调和适应保证了蝶形花亚科植物传粉过程的顺利完成。