开花时间与伴生种对鹤首马先蒿传粉和生殖成功的影响

开花物候是物种间相互作用的重要生活史特征和适合度因子,在全球气候变化的背景下而备受关注.为探讨开花时间如何存种内和种间水平上影响植物的传粉和生殖成功,我们连续3年(2003-2005)对不同花期和伴生种存在情况下的鹤首马先蒿(Pedicularis gruina)的传粉者访花忠实度、受粉率、坐果率、单果种子产量和果实被啃食频率进行了比较研究.结果表明鹤首马先蒿的坐果率主要受其传粉环境的影响:在没有伴生种时,不同时期鹤首马先蒿的坐果率没有显著差异,34-38％的花可以坐果;在有伴生种存在时既可以显著提高其坐果率,也可显著降低其坐果率,这取决于传粉者类型以及伴生种密穗马先蒿(P.densispica)花期的差异.密穗马先蒿具有花蜜和花粉双重报酬,在群落中可以作为主体物种吸引传粉者并间接促进与其伴生的鹤首马先蒿的传粉和生殖成功.同样无蜜的管花马先蒿(P.siphonantha)和鹤首马先蒿伴生,则是通过提高群落水平对传粉者的吸引力进而直接促进鹤首马先蒿的传粉和生殖成功.此外,研究结果也表明开花时间对坐果率没有显著影响,但是显著影响单果种子产量和果实被啃食的频率.在相同的传粉条件下早花期植株单果种子产量显著高于晚花期的种子产量,同时早花期的果实被啃食的频率显著增加.     

How generalists coexist: the role of floral phenotype and spatial factors in the pollination systems of two Ranunculus species

Aims Competition for pollinators between phenotypically similar flowers is believed to play an important role in floral trait diversification in the angiosperms. However, in many plant communities, species with apparently similar floral phenotypes and generalist pollination systems co-flower. Here, the pollination systems of Ranunculus acris L. and Ranunculus repens L. were investigated to determine the factors enabling the species to coexist within apparently overlapping pollination niches.Methods Sympatrically flowering populations of R. acris and R. repens were investigated at three study sites in West Wales. The floral phenotypes of the two species were compared using measurements of floral morphology and spectral analyses of petal reflectance, using principal component analysis and bee and fly colour-space models. Evidence of inter-specific discrimination by foraging insects was tested for in the field and using floral arrays. The relative roles of behavioural constancy and spatial patchiness in maintaining pollinator fidelity were estimated.Important findings The floral phenotypes of R. acris and R. repens differed significantly. Social bees were highly constant when foraging at flowers of the two species and patchy floral distribution explained some of the observed fidelity. Dipterans visiting mixed floral arrays appeared to discriminate between the species, visiting more R. acris than R. repens flowers, but there was no difference in the number of visits to single-species arrays. Social bees were more likely to display constancy to flowers of R. repens in the field. Patchiness in floral distribution, subtle differences in floral phenotype, pollinator preferences and behavioural constancy are all likely to contribute to the continued coexistence of R. acris and R. repens, despite apparent overlap in their pollination niches. Such differences have the potential to facilitate the maintenance of species diversity in plant communities, even where plants appear to share similar floral phenotypes.     

How to look like a mallow: evidence of floral mimicry between Turneraceae and Malvaceae

Abundant, many-flowered plants represent reliable and rich food sources for animal pollinators, and may even sustain guilds of specialized pollinators. Contrastingly, rare plants need alternative strategies to ensure pollinators' visitation and faithfulness. Flower mimicry, i.e. the sharing of a similar flower colour and display pattern by different plant species, is a means by which a rare species can exploit a successful model and increase its pollination services. The relationship between two or more rewarding flower mimic species, or Müllerian mimicry, has been proposed as mutualistic, in contrast to the unilaterally beneficial Batesian floral mimicry. In this work, we show that two different geographical colour phenotypes of Turnera sidoides ssp. pinnatifida resemble co-flowering Malvaceae in colour as seen by bees' eyes, and that these pollinators do not distinguish between them when approaching flowers in choice tests. Main pollinators of T. sidoides are bees specialized for collecting pollen in Malvaceae. We demonstrate that the similarity between at least one of the geographical colour phenotypes of T. sidoides and co-flowering Malvaceae is adaptive, since the former obtains more pollination services when growing together with its model than when growing alone. Instead of the convergent evolution pattern attributed to Müllerian mimicry, our data rather suggest an advergent evolution pattern, because only T. sidoides seems to have evolved to be more similar to its malvaceous models.     

The role of pollinators in floral diversification in a clade of generalist flowers

Pollinator‐mediated evolutionary divergence has seldom been explored in generalist clades because it is assumed that pollinators in those clades exert weak and conflicting selection. We investigate whether pollinators shape floral diversification in a pollination generalist plant genus, Erysimum. Species from this genus have flowers that appeal to broad assemblages of pollinators. Nevertheless, we recently reported that it is possible to sort plant species into pollination niches varying in the quantitative composition of pollinators. We test here whether floral characters of Erysimum have evolved as a consequence of shifts among pollination niches. For this, we quantified the evolutionary lability of the floral traits and their phylogenetic association with pollination niches. As with pollination niches, Erysimum floral traits show weak phylogenetic signal. Moreover, floral shape and color are phylogenetically associated with pollination niche. In particular, plants belonging to a pollination niche dominated by long‐tongued large bees have lilac corollas with parallel petals. Further analyses suggest, however, that changes in color preceded changes in pollination niche. Pollinators seem to have driven the evolution of corolla shape, whereas the association between pollination niche and corolla color has probably arisen by lilac‐flowered Erysimum moving toward certain pollination niches for other adaptive reasons.     

Functional Significance of Interspecific Variation in Burmeistera Flower Morphology: Evidence from Nectar Bat Captures in Ecuador

What causes flowers to diverge? While a plant's primary pollinator should strongly influence floral phenotype, selective pressures may also be exerted by other flower visitors or competition with other plants for pollination. Species of the primarily bat‐pollinated genus Burmeistera (Campanulaceae) frequently cooccur, with up to four species in a given site, and broadly overlap in flowering phenology, typically flowering throughout the year. The genus displays extensive interspecific variation in floral morphology in the degree that the reproductive parts (anthers and stigma) are exserted outside of the corolla, and species can be roughly classified as either long or short‐exserted. I tested two hypotheses regarding the functional significance of such variation: (1) exsertion lengths correspond to pollination by bat species of different sizes; and (2) variation serves to partition pollinator's bodies spatially and thus reduces interspecific pollen transfer. I captured bats in Ecuador to evaluate the identity and location of the Burmeistera pollen they were carrying. Results show that exsertion does not correspond to specialization on different pollinators; different bat species carried pollen of both flower types just as frequently. In support of the second hypothesis, pollen from flowers of different exsertion lengths was found to occur on different regions of bats' heads. This may serve to reduce competition for pollination among coexisting Burmeistera.     

The presence of co-flowering species facilitates reproductive success of Pedicularis monbeigiana (Orobanchaceae) through variation in bumble-bee foraging behaviour

Background and Aims The presence of co-flowering species can alter pollinator foraging behaviour and, in turn, positively or negatively affect the reproductive success of the focal species. Such interactions were investigated between a focal species, Pedicularis monbeigiana, and a co-flowering species, Vicia dichroantha, which was mediated by behaviour alteration of the shared bumble-bee pollinator. Methods Floral display size and floral colour change of P. monbeigiana were compared between pure (P. monbeigiana only) and mixed (P. monbeigiana and V. dichroantha) plots in two populations. Pollinator visitation rates, interspecific floral switching and successive within-plant pollinator visits were recorded. In addition, supplemental pollination at plant level was performed, and the fruit set and seed set were analysed in pure and mixed plots with different densities of P. monbeigiana. Key Results Pollinator visitation rates were dramatically higher in mixed plots than in pure plots. The higher pollinator visitation rates were recorded in both low- and high-density plots. In particular, successive flower visits within an individual plant were significantly lower in mixed plots. Supplemental pollination significantly increased fruit set and seed set of individuals in pure plots, while it only marginally increased seed set per fruit of plants in mixed plots. Conclusions The presence of V. dichroantha can facilitate pollination and increase female reproductive success of P. monbeigiana via both quantity (mitigating pollinator limitation) and quality (reducing geitonogamy) effects. This study suggests that successive pollinator movements among flowers within a plant, as well as pollinator visitation rates and interspecific flower switching, may be important determinants of the direction and mechanisms of interaction between species.     

Is floral morphology a good predictor of floral visitors to Antirrhineae (snapdragons and relatives)?

• The association between plants and flower visitors has been historically proposed as a main factor driving the evolutionary change of both flower and pollinator phenotypes. The considerable diversity in floral morphology within the tribe Antirrhineae has been traditionally related to pollinator types. We used empirical data on the flower visitors from 59 Antirrhineae taxa from the literature and our own field surveys, which provide an opportunity to test whether flower phenotypes are reliable predictors of visitors and pollinator niches.
• The degree of adjustment between eight key floral traits and actual visitors was explored by testing the predictive value of inferred pollinator syndromes (i.e. suites of floral traits that characterise groups of plant species related to pollination). Actual visitors and inferred pollinator niches (categorisation of visitors’ association using a modularity algorithm) were also explored using Linear Discriminant Analysis (LDA).
• The bee pollinator niche is correctly classified for flowers with dull corolla colour, without nectar guides, as the most important predictor. Both predictive value and statistical classification prove useful in classifying Antirrhineae taxa and the bee pollinator niche, mostly as a consequence of the high proportion of genera and taxa with occluded corollas primarily visited by bees. Our predictive approach rendered a high Positive Predictive Value (PPV) of floral traits in the diagnosis of visitors/pollinator niches. In particular, a high PPV was found for bees as both visitors and forming pollinator niches. In addition, LDA showed that four pollinator niches are well defined based on floral traits.
• The large number of species visited by bees irrespective of pollinator syndromes leads us to hypothesise their generalist pollinator role, despite the phenotypically specialised flowers of Antirrhineae.

     

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

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

Divergence on floral traits and vertebrate pollinators of two endemic Encholirium bromeliads

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.     

Co-flowering neighbors influence the diversity and identity of pollinator groups visiting plant species

The generalization–specialization continuum exhibited in pollination interactions currently receives much attention. It is well-known that the pollinator assemblage of particular species varies temporally and spatially, and therefore the ecological generalization on pollinators may be a contextual attribute. However, the factors causing such variation and its ecological and evolutionary consequences are still poorly understood. This variation can be caused by spatial or temporal variation in the pollinator community, but also by variation in the plant community. Here, we examined how the floral neighbourhood influenced the generalization on pollinators and the composition of pollinators of six plant species differing in generalization levels and main pollinators. The diversity, identity and density of floral species affected both the level of generalization on pollinators and the composition of visitors of particular plant species. Although the relationships to floral neighbourhood varied considerably among species, generalization level and visitation by uncommon pollinators generally increased with floral diversity and richness. The generalization level of the neighbourhood was negatively related to the generalization level of the focal species in two species. The number of flowers of the pollinator-sharing species and the number of flowers of the focal species had different effects on the composition of visits in different species; attributable to differences in facilitation/competition for pollinator attraction. We propose that an important ecological implication of our results is that variation in species interactions caused by the pollination context may result in increased community stability. The main evolutionary implication of our results is that selection on flower and pollinator traits may depend, to an unknown extent, on the composition of the co-flowering plant community.     

Pollen and stamen mimicry: the alpine flora as a case study

Many melittophilous flowers display yellow and UV-absorbing floral guides that resemble the most common colour of pollen and anthers. The yellow coloured anthers and pollen and the similarly coloured flower guides are described as key features of a pollen and stamen mimicry system. In this study, we investigated the entire angiosperm flora of the Alps with regard to visually displayed pollen and floral guides. All species were checked for the presence of pollen- and stamen-imitating structures using colour photographs. Most flowering plants of the Alps display yellow pollen and at least 28% of the species display pollen- or stamen-imitating structures. The most frequent types of pollen and stamen imitations were (mostly yellow and UV-absorbing) colour patches on petals (65% of species displaying imitations), patterns of inflorescences (18%), stamen-like pistils (10%), and staminodes (6%), as well as three-dimensional structures such as convex lower lips and filamental hairs (<5%). Dichogamous and diclinous species display pollen- and stamen-imitating structures more often than non-dichogamous and non-diclinous species, respectively. The visual similarity between the androecium and other floral organs is attributed to mimicry, i.e. deception caused by the flower visitor’s inability to discriminate between model and mimic, sensory exploitation, and signal standardisation among floral morphs, flowering phases, and co-flowering species. We critically discuss deviant pollen and stamen mimicry concepts and evaluate the frequent evolution of pollen-imitating structures in view of the conflicting use of pollen for pollination in flowering plants and provision of pollen for offspring in bees.     

Diversification of floral orientation in Lonicera is associated with pollinator shift and flowering phenology

Floral orientation has been suggested to be a functional floral trait that can enhance reproductive fitness through adaptation to both biotic factors (e.g., pollination) and abiotic factors (e.g., temperature). However, whether those factors drive the diversification of floral orientation needs to be explored through multiple related species. Phylogenetic relationships in Lonicera were identified by incorporating the species’ floral orientation (upward vs. downward) and pollinator types. Furthermore, transitions in floral orientation were evaluated together with pollinator shift, and seven Lonicera species were used to detect the influence of floral orientation on pollination and reproductive success. The relationship between temperature and the angle of floral orientation was investigated. Floral orientation in Lonicera showed that most of the closely related species within the same node displayed a similar pattern of floral orientation, and pollinator shift can lead to transitions between upward and downward orientation. Additionally, the angle of floral orientation increased with flowering phenology from spring to summer, and changing floral orientation significantly decreased pollination and seed production. Moreover, the average daily temperature during flowering phenology was significantly correlated with the angle of floral orientation. Our results indicate that pollinator shift could be an important event, leading to the diversification of floral orientation. Results of field investigations inferred that floral orientation should be a functional floral trait adapted to flowering phenology. Downward-facing flowers might help decrease heat loss to adapt to cold conditions. This study disentangled the influences of historical events and local adaptation on the evolution of floral orientation.     

Differential selection on floral traits of Ipomopsis aggregata growing in contrasting environments

Interactions for pollination between co-flowering plant species have been hypothesized to shape the evolution of their floral traits, but this hypothesis has rarely been tested. I tested the prediction that the presence of a co-flowering plant species influences the strength and/or direction of pollinator-mediated selection on floral traits. I measured phenotypic selection via female fitness on four floral traits of Ipomopsis aggregata in five populations. Three contained only conspecifics ( I only ) and two also contained the co-flowering species Penstemon barbatus ( P + I ). Directional selection via fruits/plant on corolla length and width differed in both strength and direction between P + I and I only populations. On average, selection on corolla length and width (1) was stronger in P + I than I only populations and (2) was consistently negative in P + I populations, but consistently positive in I only populations. However, these differences in selection on I . aggregata between P + I and I only populations were not caused by interactions for pollination with P . barbatus . Although plants in P + I populations received approximately 31% less conspecific pollen/flower than plants in I only populations, this difference in pollination did not translate into differences in reproductive success, which indicates that P . barbatus and I . aggregata do not strongly compete for pollination. In addition, selection via fruits/plant and conspecific pollen deposited/flower was not congruent. For example, selection on corolla length via pollen/flower was uniformly positive and did not differ between P + I and I only populations. These data suggest that the presence of P . barbatus does influence selection on floral traits of I . aggregata , but not by influencing pollination. Instead, differences in selection between P + I and I only populations appear to be the result of post-pollination modification of selection by a factor correlated with the presence of P . barbatus .     

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

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

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.     

Floral divergence, pollinator partitioning and the spatiotemporal pattern of plant–pollinator interactions in three sympatric Adenophora species

Floral divergence among congeners may relate to differential utilization of pollinators and contribute to reducing overlap in pollination niches. To investigate whether and how floral differences are associated with differential utilization of pollinators in three sympatric Adenophora species, we analyzed floral traits and evaluated the contribution of different visitors to pollination. We compared visitation rates of different pollinator categories in different years and sites. A suite of floral traits differed among the three Adenophora species, suggesting adaptation to diurnal versus nocturnal pollination and an intermediate condition. However, many visitor species were shared among the three plant species, suggesting that floral traits did not rigorously filter visitors. Effective pollinators were large bees and moths. The importance of large bees as pollinators decreased whereas that of moths increased along the gradient from typically bee-pollinated to moth-pollinated flowers. The intermediate species (A. khasiana) differed substantially from the other two species in pollinator species but not in pollinator categories. The principal pollinator category of each species was constant across years and sites except in the intermediate species where it differed between two sites. Overall, the three sympatric species of Adenophora partition pollinators by floral divergence and the principal pollinators coincide with the predictions based on floral syndromes.     

Bees explain floral variation in a recent radiation of Linaria

The role of pollinators in floral divergence has long attracted the attention of evolutionary biologists. Although abundant studies have reported the effect of pollinators on flower‐shape variation and plant speciation, the influence of pollinators on plant species differentiation during rapid radiations and the specific consequences of shifts among similar pollinators are not well understood. Here, we evaluate the association between pollinators and floral morphology in a closely related and recently diversifying clade of Linaria species (sect. Supinae subsect. Supinae). Our approach combined pollinator observations, functional floral morphometric measures and phylogenetic comparative analyses. The fauna visiting Linaria species was determined by extensive surveys and categorized by a modularity algorithm, and the size and shape of flowers were analysed by means of standard and geometric morphometric measures. Standard measures failed to find relationships between the sizes of representative pollinators and flowers. However, discriminant function analyses of geometric morphometric data revealed that pollination niches are finer predictors of flower morphologies in Linaria if compared with phylogenetic relationships. Species with the most restrictive flowers displayed the most slender spurs and were pollinated by bees with larger proboscides. These restrictive flower shapes likely appeared more than once during the evolutionary history of the study group. We show that floral variation can be driven by shifts between pollinators that have been traditionally included in a single functional group, and discuss the consequences of such transitions for plant species differentiation during rapid radiations.     

Generalized and complementary pollination system in the Andean cactus Echinopsis schickendantzii

The considerable floral diversity present in the cactus family has often been associated with the specificity of its pollinators. However, many cactus pollination systems are generalized as their flowers are pollinated by a wide spectrum of animals. For example, cactus species with white flowers, nocturnal anthesis and extended floral cycles would present generalized pollination systems in which both nocturnal and diurnal visitors could be effective pollinators. In this article, we tested this hypothesis by studying the pollination biology of Echinopsis schickendantzii, an Andean cactus with sphingophilous flowers. In addition, we evaluated whether the cactus’s pollination system is complementary or redundant regarding the relative contributions of nocturnal and diurnal pollinators. Specifically, we studied the floral cycle, the reproductive system and the pollination effectiveness of floral visitors. The flowers of E. schickendantzii are self-incompatible; they opened at crepuscule and have an extended floral cycle. Moths were frequent visitors at night, whereas bees were frequent visitors during the day; both were effective pollinators of the cactus. Our results indicated that the flowers of this species present phenotypic, functional and ecological generalization, and their fruit set is determined by the contributions of both pollinator functional groups, i.e., they have complementary pollination systems. These results support the hypothesis that cacti in the extra-tropical deserts of South America have generalized pollination systems.     

A multivariate search for pollination syndromes among penstemons

The seeming ubiquity of spatio-temporal variation in pollination regime suggests that flowers ought to be adapted to a wide range of pollinators, yet many comparative biologists perceive that in groups with complex flowers there is considerable specialization onto pollination syndromes. Statistical documentation of such syndromes has been presented for very few groups of flowers. Accordingly, we measured, for 49 species of Penstemon and close relatives, both the morphology of the flowers and visitation by pollinators. We describe the mechanics of pollination for representative species. Ordinations show a distinct difference between hummingbird-pollinated species and hymenopteran-pollinated species. Flower color is particularly good at separating hummingbird- from hymenopteran-flowers. Other characters are also correlated with this dichotomy. Within the hymenopteran-pollinated species, there are additional relationships between floral morphology and the size of the principal pollinators. Flowers frequented by large bees, such as Xylocopa , have large open vestibules and relatively short floral tubes. Flowers frequented by smaller bees, such as Osmia , have long narrow floral tubes. Unlike nectar-collecting bees, pollen-collecting bees tend to be attracted to flowers of the hummingbird syndrome. The overarching pattern was that syndrome characterizations were successful at predicting pollination by hummingbirds versus Hymenoptera, two types of animals that are profoundly different, but less successful at predicting visitation by one kind of bee versus another.     

Pollination in a patchily distributed lousewort is facilitated by presence of a co-flowering plant due to enhancement of quantity and quality of pollinator visits

Background and Aims

Plants surrounded by individuals of other co-flowering species may suffer a reproductive cost from interspecific pollen transfer (IPT). However, differences in floral architecture may reduce or eliminate IPT.

Methods

A study was made of Pedicularis densispica (lousewort) and its common co-flowering species, Astragalus pastorius, to compare reproductive and pollination success of lousewort plants from pure and mixed patches. Floral architecture and pollinator behaviour on flowers of the two plants were compared along with the composition of stigmatic pollen load of the louseworts. The extent of pollen limitation of plants from pure and mixed patches was also explored through supplemental pollination with self- and outcross pollen (PLs and PLx).

Key Results

Mixed patches attracted many more nectar-searching individuals of Bombus richardsi. These bumble-bees moved frequently between flowers of the two species. However, they pollinated P. densispica with their dorsum and A. pastorius with their abdomen. This difference in handling almost completely eliminated IPT. Lousewort plants from mixed patches yielded more seeds, and seeds of higher mass and germinability, than those from pure patches. Moreover, louseworts from mixed patches had lower PLs and PLx compared with those from pure patches.

Conclusions

Differences in floral architecture induced differences in pollinator behaviour that minimized IPT, such that co-flowering plants significantly enhanced quantity and quality of pollinator visits for the lousewort plants in patchy habitat. These findings add to our understanding of the mechanisms of pollination facilitation.