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1.
Shifts in pollen vectors favour diversification of floral traits, and differences in pollination strategies between congeneric sympatric species can contribute to reproductive isolation. Divergence in flowering phenology and selfing could also reduce interspecific crossing between self‐compatible species. We investigated floral traits and visitation rates of pollinators of two sympatric Encholirium species on rocky outcrops to evaluate whether prior knowledge of floral characters could indicate actual pollinators. Data on flowering phenology, visitation rates and breeding system were used to evaluate reproductive isolation. Flowering phenology overlapped between species, but there were differences in floral characters, nectar volume and concentration. Several hummingbird species visited flowers of both Encholirium spp., but the endemic bat Lonchophylla bokermanni and an unidentified sphingid only visited E. vogelii. Pollination treatments demonstrated that E. heloisae and E. vogelii were partially self‐compatible, with weak pollen limitation to seed set. Herbivores feeding on inflorescences decreased reproductive output of both species, but for E. vogelii the damage was higher. Our results indicate that actual pollinators can be known beforehand through floral traits, in agreement with pollination syndromes stating that a set of floral traits can be associated with the attraction of specific groups of pollinators. Divergence on floral traits and pollinator assemblage indicate that shifts in pollination strategies contribute to reproductive isolation between these Encholirium species, not divergence on flowering phenology or selfing. We suggest that hummingbird pollination might be the ancestral condition in Encholirium and that evolution of bat pollination made a substantial contribution to the diversification of this clade.  相似文献   

2.
We compared the diversity, taxonomic composition, and pollination syndromes of bromeliad assemblages and the diversity and abundance of hummingbirds along two climatically contrasting elevational gradients in Bolivia. Elevational patterns of bromeliad species richness differed noticeably between transects. Along the continuously wet Carrasco transect, species richness peaked at mid‐elevations, whereas at Masicurí most species were found in the hot, semiarid lowlands. Bromeliad assemblages were dominated by large epiphytic tank bromeliads at Carrasco and by small epiphytic, atmospheric tillandsias at Masicurí. In contrast to the epiphytic taxa, terrestrial bromeliads showed similar distributions across both transects. At Carrasco, hummingbird‐pollination was the most common pollination mode, whereas at Masicurí most species were entomophilous. The proportion of ornithophilous species increased with elevation on both transects, whereas entomophily showed the opposite pattern. At Carrasco, the percentage of ornithophilous bromeliad species was significantly correlated with hummingbird abundance but not with hummingbird species richness. Bat‐pollination was linked to humid, tropical conditions in accordance with the high species richness of bats in tropical lowlands. At Carrasco, mixed hummingbird/bat‐pollination was found especially at mid‐elevations, i.e., on the transition between preferential bat‐pollination in the lowlands and preferential hummingbird‐pollination in the highlands. In conclusion, both richness patterns and pollination syndromes of bromeliad assemblages varied in distinct and readily interpretable ways in relation to environmental humidity and temperature, and bromeliad pollination syndromes appear to follow the elevational gradients exhibited by their pollinators.  相似文献   

3.
Pollination systems frequently reflect adaptations to particular groups of pollinators. Such systems are indicative of evolutionary specialization and have been important in angiosperm diversification. We studied the evolution of pollination systems in the large genus Ruellia. Phylogenetic analyses, morphological ordinations, ancestral state reconstructions, and a character mapping simulation were conducted to reveal key patterns in the direction and lability of floral characters associated with pollination. We found significant floral morphological differences among species that were generally associated with different groups of floral visitors. Floral evolution has been highly labile and also directional. Some specialized systems such as hawkmoth or bat pollination are likely evolutionary dead-ends. In contrast, specialized pollination by hummingbirds is clearly not a dead-end. We found evidence for multiple reverse transitions from presumed ancestral hummingbird pollination to more derived bee or insect pollination. These repeated origins of insect pollination from hummingbird-pollinated ancestors have not evolved without historical baggage. Flowers of insect-pollinated species derived from hummingbird-pollinated ancestors are morphologically more similar to hummingbird flowers than they are to other more distantly related insect-pollinated flowers. Finally, some pollinator switches were concomitant with changes in floral morphology that are associated with those pollinators. These observations are consistent with the hypothesis that some transitions have been adaptive in the evolution of Ruellia.  相似文献   

4.
The flora of southern Africa has exceptional species richness and endemism, making it an ideal system for studying the patterns and processes of evolutionary diversification. Using a wealth of recent case studies, I examine the evidence for pollinator-driven diversification in this flora. Pollination systems, which represent available niches for ecological diversification, are characterized in southern Africa by a high level of ecological and evolutionary specialization on the part of plants, and, in some cases, by pollinators as well. These systems are asymmetric, with entire plant guilds commonly specialized for a particular pollinator species or functional type, resulting in obvious convergent floral evolution among guild members. Identified modes of plant lineage diversification involving adaptation to pollinators in these guilds include (i) shifts between pollination systems, (ii) divergent use of the same pollinator, (iii) coevolution, (iv) trait tracking, and (v) floral mimicry of different model species. Microevolutionary studies confirm that pollinator shifts can be precipitated when a plant species encounters a novel pollinator fauna on its range margin, and macroevolutionary studies confirm frequent pollinator shifts associated with lineage diversification. As Darwin first noted, evolutionary specialization for particular pollinators, when resulting in ecological dependency, may increase the risk of plant extinction. I thus also consider the evidence that disturbance provokes pollination failure in some southern African plants with specialized pollination systems.  相似文献   

5.
The idea of pollination syndromes has been largely discussed but no formal quantitative evaluation has yet been conducted across angiosperms. We present the first systematic review of pollination syndromes that quantitatively tests whether the most effective pollinators for a species can be inferred from suites of floral traits for 417 plant species. Our results support the syndrome concept, indicating that convergent floral evolution is driven by adaptation to the most effective pollinator group. The predictability of pollination syndromes is greater in pollinator‐dependent species and in plants from tropical regions. Many plant species also have secondary pollinators that generally correspond to the ancestral pollinators documented in evolutionary studies. We discuss the utility and limitations of pollination syndromes and the role of secondary pollinators to understand floral ecology and evolution.  相似文献   

6.
Studies of floral ecology and evolution are often centered on the idea that particular floral trait combinations, or syndromes, represent adaptations for particular pollinators. Despite the conceptual importance of pollination syndromes, few macroevolutionary studies have statistically examined the relationship between pollinators and floral traits. Using 15 species of Iochroma, Smith et al. applied phylogenetically structured correlation analyses to test the relationship between floral variation and pollination system, quantified in terms of the importance of major pollinator groups. This study revealed that pollinator shifts are tied to changes in nectar reward and floral display but are not significantly correlated with changes in corolla length or color, contrary to what might be predicted from classical pollination syndromes. Fenster et al. question these findings because our pollinator importance estimates included recently introduced honey bees. To address this concern, we recalculated importance values excluding honey bees and repeated the analyses. We found the same patterns as in our original study with significant correlations between pollinators and nectar reward and display. We conclude that phylogenetic approaches provide essential tools for testing macroevolutionary predictions of pollination syndromes and, by applying these approaches to other radiations, we can refine our understanding of the role of pollinators in floral diversification.  相似文献   

7.

Background

The hypothesis that pollinators have been important drivers of angiosperm diversity dates back to Darwin, and remains an important research topic today. Mounting evidence indicates that pollinators have the potential to drive diversification at several different stages of the evolutionary process. Microevolutionary studies have provided evidence for pollinator-mediated floral adaptation, while macroevolutionary evidence supports a general pattern of pollinator-driven diversification of angiosperms. However, the overarching issue of whether, and how, shifts in pollination system drive plant speciation represents a critical gap in knowledge. Bridging this gap is crucial to fully understand whether pollinator-driven microevolution accounts for the observed macroevolutionary patterns. Testable predictions about pollinator-driven speciation can be derived from the theory of ecological speciation, according to which adaptation (microevolution) and speciation (macroevolution) are directly linked. This theory is a particularly suitable framework for evaluating evidence for the processes underlying shifts in pollination systems and their potential consequences for the evolution of reproductive isolation and speciation.

Scope

This Viewpoint paper focuses on evidence for the four components of ecological speciation in the context of plant-pollinator interactions, namely (1) the role of pollinators as selective agents, (2) floral trait divergence, including the evolution of ‘pollination ecotypes‘, (3) the geographical context of selection on floral traits, and (4) the role of pollinators in the evolution of reproductive isolation. This Viewpoint also serves as the introduction to a Special Issue on Pollinator-Driven Speciation in Plants. The 13 papers in this Special Issue range from microevolutionary studies of ecotypes to macroevolutionary studies of historical ecological shifts, and span a wide range of geographical areas and plant families. These studies further illustrate innovative experimental approaches, and they employ modern tools in genetics and floral trait quantification. Future advances to the field require better quantification of selection through male fitness and pollinator isolation, for instance by exploiting next-generation sequencing technologies. By combining these new tools with strategically chosen study systems, and smart experimental design, we predict that examples of pollinator-driven speciation will be among the most widespread and compelling of all cases of ecological speciation.  相似文献   

8.
I present data on flower morphology, pollination, breeding system, and genetic diversity of the columnar cactus Weber-bauerocereus weberbaueri at 2500 m elevation in southwestern Peru. Weberbauerocereus weberbaueri is a self-compatible columnar cactus that is visited and pollinated by one species of rare endemic bat, Platalina genovensium, and two species of hummingbirds, Patagona gigas and Rhodopis vesper. W. weberbaueri exhibits pronounced interplant variation in floral color and size, and flowers exhibit traits corresponding to both bat and hummingbird pollination syndromes. Starch-gel electrophoresis of flower bud tissue indicated that W. weberbaueri is an autotetraploid and that genetic diversity (Hep = 0.257) of the study population was high relative to diploid plants but similar to other tetraploid species. Initial fruit set from pollinator exclusion experiments conducted in 1991 and 1993, at the onset of and after a drought associated with the El Niño event of 1991–1992, revealed that bats were the most important pollinators in 1991, but that hummingbirds and diurnal insects were most important in 1993. In both years, however, autogamy and lepidopteran larval infestation of fruits reduced differences in mature fruit production among pollinator exclusion treatments so that differences in mature fruit set were not statistically significant. Reduced bat pollination in 1993 is attributed to the reduced abundance of bats at the study site during a drought caused by El Niño. I hypothesize that interaction among several factors, including tetraploidy, autogamy, larval infestation of developing fruits, and variation in pollinator abundance, may not result in strong selection for a bat vs. hummingbird floral morph, thus allowing the persistence of floral variation in this cactus.  相似文献   

9.
Covariation among traits can modify the evolutionary trajectory of complex structures. This process is thought to operate at a microevolutionary scale, but its long‐term effects remain controversial because trait covariation can itself evolve. Flower morphology, and particularly floral trait (co)variation, has been envisioned as the product of pollinator‐mediated selection. Available evidence suggests that major changes in pollinator assemblages may affect the joint expression of floral traits and their phenotypic integration. We expect species within a monophyletic lineage sharing the same pollinator type will show not only similarity in trait means but also similar phenotypic variance‐covariance structures. Here, we tested this expectation using eighteen Salvia species pollinated either by bees or by hummingbirds. Our findings indicated a nonsignificant multivariate phylogenetic signal and a decoupling between means and variance‐covariance phenotypic matrices of floral traits during the evolution to hummingbird pollination. Mean trait value analyses revealed significant differences between bee‐ and hummingbird‐pollinated Salvia species although fewer differences were detected in the covariance structure between groups. Variance‐covariance matrices were much more similar among bee‐ than hummingbird‐pollinated species. This pattern is consistent with the expectation that, unlike hummingbirds, bees physically manipulate the flower, presumably exerting stronger selection pressures favouring morphological convergence among species. Overall, we conclude that the evolution of hummingbird pollination proceeded through different independent transitions. Thus, although the evolution of hummingbird pollination led to a new phenotypic optimum, the process involved the diversification of the covariance structure.  相似文献   

10.
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.  相似文献   

11.
兰科植物欺骗性传粉   总被引:7,自引:0,他引:7  
植物与传粉动物的互利关系在生态系统中非常普遍。然而,有许多植物不为传粉者提供任何报酬,而是利用各种欺骗方式诱骗昆虫拜访,从而实现传粉,即欺骗性传粉。兰科是被子植物大科之一,其高度特化的繁殖器官和适应于昆虫传粉的精巧结构令人称奇。进化论创始人达尔文描述了许多兰花与昆虫精巧的传粉系统,但他忽视了欺骗性传粉的存在。事实上,近1/3的兰科植物都依赖于欺骗性传粉。欺骗性传粉可能是导致兰科植物多样性的重要原因之一。兰花利用或操作昆虫觅食、交配、产卵和栖息等行为,演化出各种各样的欺骗性传粉机制,常见的类型包括泛化的食源性欺骗、Batesian拟态、性欺骗、产卵地拟态和栖息地拟态。花的颜色、形态和气味在欺骗性传粉的成功实现中起到了重要作用。欺骗性兰花与传粉昆虫之间的演化可能是不同步的,兰花追踪昆虫的行为信号而发生分化,然而欺骗性传粉可能对昆虫造成一定的伤害,从而对昆虫也施加选择压力。由于昆虫的学习行为,欺骗性的兰花一般具有低的昆虫拜访率和结实率,其繁殖成功率受各种因素的影响。欺骗性加剧了兰花对传粉昆虫的依赖,使其具有更高的灭绝风险,传粉生物学的研究能为兰科植物的有效保护提供指导。在欺骗性传粉系统中,有报酬的伴生植物、拟态模型和其他拟态信号提供者对传粉成功有重要影响。因此,研究欺骗性传粉兰花、传粉昆虫和相关的生物和生态因子的网状进化关系具有重要理论和实践意义。  相似文献   

12.
Flowers show important structural variation as reproductive organs but the evolutionary forces underlying this diversity are still poorly understood. In animal‐pollinated species, flower shape is strongly fashioned by selection imposed by pollinators, which is expected to vary according to guilds of effective pollinators. Using the Antillean subtribe Gesneriinae (Gesneriaceae), we tested the hypothesis that pollination specialists pollinated by one functional type of pollinator have maintained more similar corolla shapes through time due to more constant and stronger selection constraints compared to species with more generalist pollination strategies. Using geometric morphometrics and evolutionary models, we showed that the corolla of hummingbird specialists, bat specialists, and species with a mixed‐pollination strategy (pollinated by hummingbirds and bats; thus a more generalist strategy) have distinct shapes and that these shapes have evolved under evolutionary constraints. However, we did not find support for greater disparity in corolla shape of more generalist species. This could be because the corolla shape of more generalist species in subtribe Gesneriinae, which has evolved multiple times, is finely adapted to be effectively pollinated by both bats and hummingbirds. These results suggest that ecological generalization is not necessarily associated with relaxed selection constraints.  相似文献   

13.
Concerted changes in flower morphology and pollinators provide strong evidence on adaptive evolution. Schizanthus (Solanaceae) has zygomorphic flowers and consists of 12 species of annual or biennial herbs that are distributed mainly in Chile and characterized by bee-, hummingbird-, and moth-pollination syndromes. To infer whether flowers diversified in relation to pollinator shifts, we traced the evolutionary trajectory of flower traits and visitors onto a phylogeny based on sequence data from ITS, waxy, and trnF/ndhJ DNA. Maximum-likelihood ancestral reconstruction of floral traits suggests that ancestral Schizanthus had a bee-pollination syndrome. The hummingbird syndrome evolved in S. grahamii, a high elevation species in the Andes. The moth syndrome evolved in the ancestor of three species that inhabit the Atacama Desert. Results of mapping flower visitors onto the phylogeny show that the shift from bee to hummingbird pollination concurred with a shift in pollinators as predicted by the syndromes. However, the same pattern was not found for the moth syndrome. Visits by moths were observed only in one of the three moth-syndrome species, and at a very low rate. This mismatch suggests either anachronic floral characters or maintenance of rare, imperceptible moth pollination backed up by capacity for autonomous selfing. Overall, results suggest that diversification of flower traits in Schizanthus has occurred in relation to pollinator shifts.  相似文献   

14.
  1. Pollination syndromes refer to stereotyped floral characteristics (flower colour, shape, etc.) that are associated with a functional group of pollinators (bee, bird, etc.).
  2. The trumpet creeper Campsis radicans, endemic to the southeast and mid‐west United States, has been assigned to the hummingbird‐pollination syndrome, due mainly to its red, trumpet‐shaped flowers.
  3. Previous studies demonstrated that the ruby‐throated hummingbird Archilochus colubris is C. radicans' primary pollinator, but anecdotal data suggest various bee species may provide pollination service when hummingbirds are absent.
  4. This study characterised C. radicans nectar volume and concentration by time of day. Nectar volume was suitable for hummingbirds, but concentration was higher than typical hummingbird‐pollinated plants (~20% w/w); at ~30% w/w, it approached the concentration expected in bee‐pollinated plants (~50% w/w). We also found substantial amounts of nectar at night.
  5. Two C. radicans populations received virtually no hummingbird visits, but the number of bees were markedly higher than in the populations previously described. Interestingly, there were no night‐time visitors despite the large quantity of nocturnal nectar.
  6. Based on previously published pollen delivery per visit by various species, this study estimated that cumulative deposition by bees routinely reached pollen deposition thresholds for setting fruit in C. radicans. They are, unequivocally, the predominant pollinators in these populations, thus providing pollination service in the absence of hummingbirds.
  7. These results highlight C. radicans as a food source for native bees and add to the understanding of how floral phenotypes can facilitate pollination by disparate functional groups.
  相似文献   

15.
The geographical distribution of pollinators is an important factor determining the biogeographical pattern of floral evolution and diversification. In Africa, a guild of plant species has converged in a floral syndrome for pollination by long‐tongued hawkmoths (predominantly Agrius convolvuli). It is hypothesized that such floral convergence could track the geographical distribution of long‐tongued hawkmoths, so it may not be confined to Africa. We investigated the pollination biology of Lilium sargentiae E. H. Wilson, which is endemic to China and exhibits traits suggestive of long‐tongued hawkmoth pollination. Lilium sargentiae was visited by A. convolvuli as well as small beetles (Nitidulidae) but pollination was mostly or totally effected by the moth. It was consistent with other hawkmoth‐pollinated plants in terms of floral tube length, nectar traits, tepal reflectance, and scent composition. We present the first experimental evidence for the hypothesis proposed above and for hawkmoth pollination in the widespread and ornamentally and economically important genus Lilium L. in a natural habitat. Our findings imply that long‐tongued hawkmoths (especially the extremely widespread A. convolvuli) as a distinctive pollinator niche may have underlain the evolution of long‐tubed (>8 cm) flowers across different continents.  相似文献   

16.
17.
18.
BACKGROUND AND AIMS: Based on the bell shape and greenish colour of the flowers, bat-pollination was suggested for some Sinningieae species (Gesneriaceae). Actually, there are no reports on pollination biology and visitors for these species. This paper reports on pollination biology of Sinningia brasiliensis, Paliavana prasinata and P. sericiflora in south-eastern Brazil. METHODS: Flowers were observed in situ to determine phases of anthesis, colour patterns and scent intensity. Corolla measures were taken from fresh flowers. Nectar production and concentration were measured in flowers bagged at the pre-anthesis phases. Direct visual observations of visitors were made during the day and night, and photographs were taken to analyse their visiting behaviour. KEY RESULTS: Some floral features of the three species fit the bat-pollination syndrome: large, robust and gullet-shaped corollas, colour patterns and large amount of nectar. However, other floral features-such as onset of anthesis, nectar attributes and odour intensity-differ among the species. Nectar volume and total sugar production increased significantly at midnight in S. brasiliensis and P. prasinata, but in P. sericiflora there were no significant differences in the total nectar and sugar production during anthesis. Scent intensity is much higher in S. brasiliensis and P. prasinata than P. sericiflora. Flowers of S. brasiliensis and P. prasinata, whose features fit the chiropterophilous syndrome, are pollinated by glossophagine bat species. Paliavana sericiflora, on the other hand, presents floral features intermediate between bat and hummingbird syndromes, but is visited and pollinated only by hummingbirds. CONCLUSIONS: These data strengthen the statement that the bat syndrome in Sinningieae originated independently in Sinningia brasiliensis and in Paliavana species. Paliavana sericiflora may be another example of a plant 'in transition' from the hummingbird to the bat pollination, but a reversion in the direction of bat to hummingbird might not be excluded. It is also possible that this is a case of speciation on both bat and hummingbird pollination. Studies on Paliavana sister species may provide insights about origins and the evolutionary directions of the pollination systems of these species.  相似文献   

19.
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.  相似文献   

20.
  • Unrelated plants adapted to particular pollinator types tend to exhibit convergent evolution in floral traits. However, inferences about likely pollinators from ‘pollination syndromes’ can be problematic due to trait overlap among some syndromes and unusual floral architecture in some lineages. An example is the rare South African parasitic plant Mystropetalon thomii (Mystropetalaceae), which has highly unusual brush‐like inflorescences that exhibit features of both bird and rodent pollination syndromes.
  • We used camera traps to record flower visitors, quantified floral spectral reflectance and nectar and scent production, experimentally determined self‐compatibility and breeding system, and studied pollen dispersal using fluorescent dyes.
  • The dark‐red inflorescences are usually monoecious, with female flowers maturing before male flowers, but some inflorescences are purely female (gynoecious). Inflorescences were visited intensively by several rodent species that carried large pollen loads, while visits by birds were extremely rare. Rodents prefer male‐ over female‐phase inflorescences, likely because of the male flowers’ higher nectar and scent production. The floral scent contains several compounds known to attract rodents. Despite the obvious pollen transfer by rodents, we found that flowers on both monoecious and gynoecious inflorescences readily set seed in the absence of rodents and even when all flower visitors are excluded.
  • Our findings suggest that seed production occurs at least partially through apomixis and that M. thomii is not ecologically dependent on its rodent pollinators. Our study adds another species and family to the growing list of rodent‐pollinated plants, thus contributing to our understanding of the floral traits associated with pollination by non‐flying mammals.
  相似文献   

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