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.     

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.     

Pollination syndromes in a specialised plant‐pollinator interaction: does floral morphology predict pollinators in Calceolaria?

Pollination syndromes are defined as suites of floral traits evolved in response to selection imposed by a particular group of pollinators (e.g., butterflies, hummingbirds, bats). Although numerous studies demonstrated their occurrence in plants pollinated by radically different pollinators, it is less known whether it is possible to identify them within species pollinated by one functional pollinator group. In such a framework, we expect floral traits to evolve also in response to pollinator subgroups (e.g., species, genera) within that unique functional group. On this, specialised pollination systems represent appropriate case studies to test such expectations. Calceolaria is a highly diversified plant genus pollinated by oil‐collecting bees in genera Centris and Chalepogenus. Variation in floral traits in Calceolaria has recently been suggested to reflect adaptations to pollinator types. However, to date no study has explicitly tested that observation. In this paper, we quantitatively test that hypothesis by evaluating the presence of pollination syndromes within the specialised pollination system formed by several Calceolaria and their insect pollinators. To do so, we use multivariate approaches and explore the structural matching between the morphology of 10 Calceolaria taxa and that of their principal pollinators. Our results identify morphological matching between floral traits related to access to the reward and insect traits involved in oil collection, confirming the presence of pollinator syndromes in Calceolaria. From a general perspective, our findings indicate that the pollination syndrome concept can be also extended to the intra‐pollinator group level.     

CONVERGENT AND CORRELATED EVOLUTION OF MAJOR LIFE‐HISTORY TRAITS IN THE ANGIOSPERM GENUS LEUCADENDRON (PROTEACEAE)

Natural selection is expected to cause convergence of life histories among taxa as well as correlated evolution of different life‐history traits. Here, we quantify the extent of convergence of five key life‐history traits (adult fire survival, seed storage, degree of sexual dimorphism, pollination mode, and seed‐dispersal mode) and test hypotheses about their correlated evolution in the genus Leucadendron (Proteaceae) from the fire‐prone South African fynbos. We reconstructed a new molecular phylogeny of this highly diverse genus that involves more taxa and molecular markers than previously. This reconstruction identifies new clades that were not detected by previous molecular study and morphological classifications. Using this new phylogeny and robust methods that account for phylogenetic uncertainty, we show that the five life‐history traits studied were labile during the evolutionary history of the genus. This diversity allowed us to tackle major questions about the correlated evolution of life‐history strategies. We found that species with longer seed‐dispersal distances tended to evolve lower pollen‐dispersal distance, that insect‐pollinated species evolved decreased sexual dimorphism, and that species with a persistent soil seed‐bank evolved toward reduced fire‐survival ability of adults.     

Insect and wind pollination of an alpine biennial Aconitum gymnandrum (Ranunculaceae)

Wind pollination can provide reproductive insurance for animal‐pollinated dioecious plants in the absence of available pollinators, but combinations of insect and wind pollination (ambophily) have rarely been studied in hermaphrodite herbs. We examined the stable occurrence of insect pollination and wind pollination over 4 years in a population of a biennial Aconitum species (A. gymnandrum) with actinomorphic and degenerate sepals. The total frequency of visits of two bumblebee species showed no distinct fluctuations in the studied population among the 4 years. However, seed production of netted flowers after emasculation indicated wind pollination had occurred. The seed number of bagged flowers with one visit by bumblebees was significantly less than that of netted flowers after one visit, or in control flowers. Both seed number and fruit set of netted flowers were significantly lower than in control flowers. These results suggest that wind pollination provides supplementary pollen to unvisited and/or once‐visited flowers, but accounts for only a small amount of seed production compared to bumblebee pollination in natural conditions. Such a combination of insect and wind pollination might play an important role in maintaining sexual reproduction of this biennial herb, allowing it to persist in arid habitats on the Qinghai‐Tibetan Plateau, especially during Quaternary glacial periods when pollinator populations oscillated extensively.     

Evolutionary transition between bee pollination and hummingbird pollination in Salvia: Comparing means,variances and covariances of corolla traits

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.     

The design of trapping devices in pollination traps of the genus Arum (Araceae) is related to insect type

Pollinators have long been known to select for floral traits, but the nature of this relationship has been little investigated in trap pollination systems. We investigated the trapping devices of 15 Arum spp. and compared them with the types of insects trapped. Most species shared a similar general design of trap chamber walls covered in downward‐pointing papillate cells, lacunose cells in the chamber wall and elongated sterile flowers partially blocking the exit of the trap. However, there was significant variation in all these morphological features between species. Furthermore, these differences related to the type of pollinator trapped. Most strikingly, species pollinated by midges had a slippery epidermal surface consisting of smaller papillae than in species pollinated by other insects. Midge‐pollinated species also had more elongated sterile flowers and tended to have a larger lacunose area. We conclude that pollination traps evolve in response to the type of insect trapped and that changes to the slippery surfaces of the chamber wall are an important and previously little recognized variable in the design of pollination traps. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 385–397.     

Floral specialization for different pollinators and divergent use of the same pollinator among co‐occurring Impatiens species (Balsaminaceae) from Southeast Asia

Floral variation among closely related species is thought to often reflect differences in pollination systems. Flowers of the large genus Impatiens are characterized by extensive variation in colour, shape and size and in anther and stigma positioning, but studies of their pollination ecology are scarce and most lack a comparative context. Consequently, the function of floral diversity in Impatiens remains enigmatic. This study documents floral variation and pollination of seven co‐occurring Impatiens spp. in the Southeast Asian diversity hotspot. To assess whether floral trait variation reflects specialization for different pollination systems, we tested whether species depend on pollinators for reproduction, identified animals that visit flowers, determined whether these visitors play a role in pollination and quantified and compared key floral traits, including floral dimensions and nectar characteristics. Experimental exclusion of insects decreased fruit and seed set significantly for all species except I. muscicola, which also received almost no visits from animals. Most species received visits from several animals, including bees, birds, butterflies and hawkmoths, only a subset of which were effective pollinators. Impatiens psittacina, I. kerriae, I. racemosa and I. daraneenae were pollinated by bees, primarily Bombus haemorrhoidalis. Impatiens chiangdaoensis and I. santisukii had bimodal pollination systems which combined bee and lepidopteran pollination. Floral traits differed significantly among species with different pollination systems. Autogamous flowers were small and spurless, and did not produce nectar; bee‐pollinated flowers had short spurs and large floral chambers with a wide entrance; and bimodally bee‐ and lepidopteran‐pollinated species had long spurs and a small floral chamber with a narrow entrance. Nectar‐producing species with different pollination systems did not differ in nectar volume and sugar concentration. Despite the high frequency of bee pollination in co‐occurring species, individuals with a morphology suggestive of hybrid origin were rare. Variation in floral architecture, including various forms of corolla asymmetry, facilitates distinct, species‐specific pollen‐placement on visiting bees. Our results show that floral morphological diversity among Impatiens spp. is associated with both differences in functional pollinator groups and divergent use of the same pollinator. Non‐homologous mechanisms of floral asymmetry are consistent with repeated independent evolution, suggesting that competitive interactions among species with the same pollination system have been an important driver of floral variation among Impatiens spp.     

Phenotypic selection on flowering phenology and size in two dioecious plant species with different pollen vectors

Dioecious plants may be pollinated biotically by animals or abiotically via wind or water currents. It has been hypothesized that these two types of pollen vectors might impose contrasting selective pressures on plant flowering phenology. In the present study we describe the flowering phenology of two sympatric dioecious species with contrasting pollination modes: Mercurialis perennis (wind‐pollinated) and Tamus communis (insect‐pollinated). We estimated selection differentials and gradients for flowering time and flowering synchrony. As flowering time might depend on the accumulation of enough internal resources, we also estimated direct and indirect selection on plant size. Both species have male‐biased sexual ratios, and males are bigger and produce larger flower displays than females, but only in T. communis do males bloom earlier and for longer than females. Selection gradients suggest that selection tends to favor early‐flowering females of T. communis. There is no evidence of direct current selection on the flowering phenology of M. perennis. Intersexual differences in phenology fit with sex allocation and sexual selection theories. As we hypothesized, phenology of the animal‐pollinated species is under stronger selection than that of the wind‐pollinated species and we discuss the potential role of pollen vectors in shaping the flowering phenologies of the study species.     

Does competition with wind‐pollinated species alter Echium plantagineum's attractiveness to a common pollinator Bombus terrestris?

1. In insect‐pollinated plants, pollinator attraction is influenced by flowers (e.g. number, size) and their associated rewards (e.g. pollen, nectar). These traits can depend on plant interactions. Indeed, below‐ground competition between plants can lead to a decrease in flower or reward production in insect‐pollinated species. 2. Wind‐pollinated plants, in particular, which are almost never studied in plant–pollinator networks, can alter insect‐pollinated plants' attractiveness through competition for nutrients. The response of pollinators to such changes has never been investigated. 3. A pot experiment was carried out in which an insect‐pollinated species, Echium plantagineum, was grown in binary mixture with three wind‐pollinated species selected to exert a panel of competitive interactions. Below‐ground competition was controlled using dividers limiting interspecific root competition. Floral traits of E. plantagineum (i.e. flower production, floral display size, flower size and nectar production) were measured. For each species mixture, the visits (i.e. first visit, number of visits, 10‐min sequences) of Bombus terrestris individuals released in a flight cage containing two pots were followed, one with and one without below‐ground competition. 4. Below‐ground competition significantly affected nectar's sucrose concentration but did not influence flower and nectar production. Likewise, pollinator visits were not influenced by below‐ground competition. Competitor identity significantly influenced flower and reward production of E. plantagineum, with a decrease in the presence of the most competitive wind‐pollinated species. A tendency for faster flower visitation events was also detected in the presence of the least competitive competitor. This study raises new questions regarding the influence of wind‐pollinated plants on plant–pollinator interactions.     

Temporal,but not spatial,changes in expression patterns of petal identity genes are associated with loss of papillate conical cells and the shift to bird pollination in Macaronesian Lotus (Leguminosae)

• In the generally bee‐pollinated genus Lotus a group of four species have evolved bird‐pollinated flowers. The floral changes in these species include altered petal orientation, shape and texture. In Lotus these characters are associated with dorsiventral petal identity, suggesting that shifts in the expression of dorsal identity genes may be involved in the evolution of bird pollination. Of particular interest is Lotus japonicus CYCLOIDEA 2 (LjCYC2), known to determine the presence of papillate conical cells on the dorsal petal in L. japonicus. Bird‐pollinated species are unusual in not having papillate conical cells on the dorsal petal.
• Using RT‐PCR at various stages of flower development, we determined the timing of expression in all petal types for the three putative petal identity genes (CYC‐like genes) in different species with contrasting floral morphology and pollination syndromes.
• In bird‐pollinated species the dorsal identity gene, LjCYC2, is not expressed at the floral stage when papillate conical cells are normally differentiating in bee‐pollinated species. In contrast, in bee‐pollinated species, LjCYC2 is expressed during conical cell development.
• Changes in the timing of expression of the above two genes are associated with modifications in petal growth and lateralisation of the dorsal and ventral petals in the bird‐pollinated species. This study indicates that changes in the timing, rather than spatial distribution, of expression likely contribute to the modifications of petal micromorphology and petal size during the transition from bee to bird pollination in Macaronesian Lotus species.

     

Experimental evidence for specialized bird pollination in the endangered South African orchid Satyrium rhodanthum and analysis of associated floral traits

The bird pollination syndrome is characterized by red, unscented flowers with dilute nectar in long nectar tubes. However, the extent to which plants with such traits actually depend on birds for seed production is seldom determined experimentally, and traits such as colour and scent production are often assessed only subjectively. We documented bird pollination and quantified floral traits in the critically endangered Satyrium rhodanthum (Orchidaceae) from mistbelt grasslands in the summer‐rainfall region of South Africa. Direct observations and motion trigger camera footage revealed amethyst sunbirds as the only pollinators, despite the presence of other potential pollinators. Experimental exclusion of sunbirds significantly reduced pollination and fruit set to near zero. Pollination success in naturally pollinated plants was close to 100% in one year, and fruit set varied from 23 to 64% in other years. Pollen transfer efficiency was 5.8%, which is lower than in related insect‐pollinated species, probably due to a tendency of birds to wipe pollinaria from their beak. Flowers of S. rhodanthum only reflect light in the red range of the spectrum, and they produce only a few aliphatic and monoterpene scent compounds at comparatively low emission rates. Nectar volume and sugar concentration varied between 2.7 and 3.7 μL and 23.7 and 25.9%, respectively. We conclude that S. rhodanthum is highly specialized for pollination by sunbirds. Colour, scent and nectar characteristics differ from insect‐pollinated Satyrium species and are consistent with those expected for bird‐pollinated flowers, and may contribute to lack of visitation by other potential long‐tongued pollinators. Habitat loss probably underlies the critically endangered conservation status of S. rhodanthum, but the specialization for pollination by a single bird species means that reproduction in this orchid is vulnerable to losses in surrounding communities of plants that subsidize the energetic requirements of sunbirds. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 177 , 141–150.     

Geographic variation of reproductive traits and competition for pollinators in a bird‐pollinated plant

Geographic variation in the reproductive traits of animal‐pollinated plants can be shaped by spatially variable selection imposed by differences in the local pollination environment. We investigated this process in Babiana ringens (Iridaceae), an enigmatic species from the Western Cape region of South Africa. B. ringens has evolved a specialized perch facilitating cross‐pollination by sunbirds and displays striking geographic variation in perch size and floral traits. Here, we investigate whether this variation can be explained by geographic differences in the pollinator communities. We measured floral and inflorescence traits, and abiotic variables (N, P, C, and rainfall) and made observations of sunbirds in populations spanning the range of B. ringens. In each population, we recorded sunbird species identity and measured visitation rates, interfloral pollen transfer, and whether the seed set of flowers was pollen limited. To evaluate whether competition from co‐occurring sunbird‐pollinated species might reduce visitation, we quantified nectar rewards in B. ringens and of other co‐flowering bird‐pollinated species in local communities in which populations occurred. Variation in abiotic variables was not associated with geographical variation of traits in B. ringens. Malachite sunbirds were the dominant visitor (97% of visits) and populations with larger‐sized traits exhibited higher visitation rates, more between‐flower pollen transfer and set more seed. No sunbirds were observed in four populations, all with smaller‐sized traits. Sunbird visitation to B. ringens was not associated with local sunbird activity in communities, but sunbird visitation was negatively associated with the amount of B. ringens sugar relative to the availability of alternative nectar sources. Our study provides evidence that B. ringens populations with larger floral traits are visited more frequently by sunbirds, and we propose that visitation rates to B. ringens may be influenced, in part, by competition with other sunbird‐pollinated species.     

The best of two worlds: ecology and evolution of ambophilous plants

Ambophily, the mixed mode of wind and insect pollination is still poorly understood, even though it has been known to science for over 130 years. While its presence has been repeatedly inferred, experimental data remain regrettably rare. No specific suite of morphological or ecological characteristics has yet been identified for ambophilous plants and their ecology and evolution remain uncertain. In this review we summarise and evaluate our current understanding of ambophily, primarily based on experimental studies. A total of 128 ambophilous species – including several agriculturally important crops – have been reported from most major habitat types worldwide, but this probably represents only a small subset of ambophilous species. Ambophilous species have evolved both from wind- and insect-pollinated ancestors, with insect-pollinated ancestors mostly representing pollination by small, generalist flower visitors. We compiled floral and reproductive traits for known ambophilous species and compared our results to traits of species pollinated either by wind or by small generalist insects only. Floral traits were found to be heterogeneous and strongly overlap especially with those of species pollinated by small generalist insects, which are also the prominent pollinator group for ambophilous plants. A few ambophilous species are only pollinated by specialised bees or beetles in addition to pollination by wind. The heterogeneity of floral traits and high similarity to generalist small insect-pollinated species lead us to conclude that ambophily is not a separate pollination syndrome but includes species belonging to different insect- as well as wind-pollination syndromes. Ambophily therefore should be regarded as a pollination mode. We found that a number of ecological factors promoted the evolution of ambophily, including avoidance of pollen limitation and self-pollination, spatial flower interference and population density. However, the individual ecological factors favouring the transition to ambophily vary among species depending on species distribution, habitat, population structure and reproductive system. Finally, a number of experimental studies in combination with observations of floral traits of living and fossil species and dated phylogenies may indicate evolutionary stability. In some clades ambophily has likely prevailed for millions of years, for example in the castanoid clade of the Fagaceae.     

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.     

Airborne conifer pollen grains are rarely deposited on stigmas of coflowering insect‐pollinated angiosperms

Although plant species with either animal or wind pollination modes are widespread and usually sympatric in nature, the degree of pollen interference from wind‐pollinated species on animal‐pollinated species remains little known. Conifer trees generally release a huge number of pollen grains into the air, floating into our noses and sometimes causing an allergic response. Here we document airborne pollen from two conifers (Pinus densata Mast. and Picea likiangensis (Franch.) E. Pritz.) deposited on the stigmas of eight coflowering insect‐pollinated angiosperms over 2 years in a mountainous forest community, in Shangri‐La, southwest China. Pollen density in the air as well as conifer pollen deposited onto stigmas at short and long distances from the airborne pollen source were quantified. Our results showed that conifer pollen as a proportion of total stigmatic pollen loads in the insect‐pollinated plants varied from 0.16% to 8.67% (3.16% ± 0.41%, n = 735) in 2016 and 0.66% to 5.38% (2.87% ± 0.86%, n = 180), and pollen quantity per unit area was closely related to that of airborne pollen in the air. Conifer pollen deposition on stigmas of insect‐pollinated species decreased greatly with increased distance from the pollen source. In the 10 plant species flowering in summer after conifer pollen release had finished, heterospecific pollen deposited on these stigmas came mainly from other insect‐pollinated flowers, with little contribution from airborne conifer pollen. The results indicate that there might be little interference with coflowering angiosperms by airborne pollen from dominant conifers in natural communities.     

The evolution of polymorphic sexual systems in daffodils (Narcissus)

Narcissus, the daffodil genus, exhibits an unusual diversity of sexual systems, with populations that are monomorphic, dimorphic or trimorphic for style length. Associated with this variation are striking differences among species in floral morphology and pollination biology. This diversity provides an opportunity to investigate the evolution of mating polymorphisms, and to determine how floral morphology promotes transitions among sexual systems. Because of the absence of heteromorphic incompatibility in Narcissus, floral morphology plays a key role in governing patterns of outcrossed mating. Phylogenetic evidence indicates that stylar monomorphism is ancestral in the genus, with multiple origins of stylar polymorphism, including independent origins of stigma-height dimorphism, distyly and tristyly. Sexual polymorphisms have evolved only in lineages with narrow floral tubes that are pollinated by Lepidoptera and/or long-tongued bees. Populations of polymorphic Narcissus species are typically dominated by the long-styled morph and display imperfect reciprocity in the positions of sexual organs. These features are consequences of the unusual association between stylar polymorphism and a self-incompatibility system that permits intramorph mating.     

Does the morphological fit between flowers and pollinators affect pollen deposition? An experimental test in a buzz‐pollinated species with anther dimorphism

Some pollination systems, such as buzz‐pollination, are associated with floral morphologies that require a close physical interaction between floral sexual organs and insect visitors. In these systems, a pollinator's size relative to the flower may be an important feature determining whether the visitor touches both male and female sexual organs and thus transfers pollen between plants efficiently. To date, few studies have addressed whether in fact the “fit” between flower and pollinator influences pollen transfer, particularly among buzz‐pollinated species. Here we use Solanum rostratum, a buzz‐pollinated plant with dimorphic anthers and mirror‐image flowers, to investigate whether the morphological fit between the pollinator's body and floral morphology influences pollen deposition. We hypothesized that when the size of the pollinator matches the separation between the sexual organs in a flower, more pollen should be transferred to the stigma than when the visitor is either too small or too big relative to the flower. To test this hypothesis, we exposed flowers of S. rostratum with varying levels of separation between sexual organs, to bumblebees (Bombus terrestris) of different sizes. We recorded the number of visits received, pollen deposition, and fruit and seed production. We found higher pollen deposition when bees were the same size or bigger than the separation between anther and stigma within a flower. We found a similar, but not statistically significant pattern for fruit set. In contrast, seed set was more likely to occur when the size of the flower exceeded the size of the bee, suggesting that other postpollination processes may be important in translating pollen receipt to seed set. Our results suggest that the fit between flower and pollinator significantly influences pollen deposition in this buzz‐pollinated species. We speculate that in buzz‐pollinated species where floral morphology and pollinators interact closely, variation in the visitor's size may determine whether it acts mainly as a pollinator or as a pollen thief (i.e., removing pollen rewards but contributing little to pollen deposition and fertilization).     

Pollen-Ovule Ratio and Gamete Investment in Pedicularis （Orobanchaceae）

The Pedicularis species provides ideal materials to study floral evolution because of their substantial flower variation based on a narrow genetic basis, even though they are almost exclusively pollinated by bumblebee. These traits allow us to detect the evolutionary trends of floral parameters without considering genetic background and the difference of pollination vectors. The pollen-ovule ratio is widely used to estimate the pattern of resource investment in two sexual functions in flowering plants. Forty species representing all of the corolla types in Pedicularls were used to study pollen-ovule ratio, gamete investment, and their correlations. Results show that pollen-ovule ratio does not differ among both different corolla types and taxonomic groups. It is therefore suggested that pollen-ovule ratio should be a parallel evolution. The correlations between pollen-ovule ratio and pollen size （-）, and ovule size （＋） can be successfully explained in terms of sex allocation theory. The biological significance of such relationships was also discussed. Additionally, we analyzed the pattern of resource investment into female gamete, which has been somewhat neglected, and found that plants have different patterns of gamete investment between the two sexual functions.     

Pollinator‐mediated selection in a specialized pollination system: matches and mismatches across populations

Most studies on pollinator‐mediated selection have been performed in generalized rather than specialized pollination systems. This situation has impeded evaluation of the extent to which selection acts on attraction or specialized key floral traits involved in the plant‐pollinator phenotypic interphase. We studied pollinator‐mediated selection in four populations of Nierembergia linariifolia, a self‐incompatible and oil‐secreting plant pollinated exclusively by oil‐collecting bees. We evaluated whether floral traits experience variable selection among populations and whether attraction and fit traits are heterogeneously selected across populations. Populations differed in every flower trait and selection was consistently observed for corolla size and flower shape, two traits involved in the first steps of the pollination process. However, we found no selection acting on mechanical‐fit traits. The observation that selection occurred upon attraction rather than mechanical‐fit traits, suggests that plants are not currently evolving fine‐tuned morphological adaptations to local pollinators and that phenotypic matching is not necessarily an expected outcome in this specialized pollination system.