Why be a rewarding trap? The evolution of floral rewards in Arum (Araceae), a genus characterized by saprophilous pollination systems

Plant species of the genus Arum typically have lure-and-trap pollination systems that are saprophilous (i.e. attracting flies or beetles searching for breeding sites in decaying organic matter). They have been assumed to always attract and trap their pollinators by deception because the inflorescences provide unsuitable breeding grounds for pollinators. The present study explored the possibility that one species, Arum creticum Boiss. & Heldr., which has yellow, sweet-smelling inflorescences, rewards its pollinators and that this increases its success in attracting pollinators over its close relative, Arum idaeum Coust. & Gadoger. The relationship between rewards provided, floral structure, insect attraction, and pollen import and export was examined in two naturally occurring sympatric populations of A. creticum , A. idaeum , and their natural hybrids. The results showed that plants providing more pollen were visited by larger numbers of females of a mining bee Lasioglossum marginatum Brullé as well as adults and nymphs of a hemipteran bug Dionconotus cruentatus creticus Heiss. In A. creticum , L. marginatum was found to be a better outcrossing vector than D. cruentatus in areas where both pollinators occurred because L. marginatum individuals carried more pollen on their bodies and travelled greater distances between inflorescences, thus increasing the potential for outcrossing. The finding that floral rewards may result in increased fitness, compared to nonrewarding systems that rely on attracting saprophilic pollinators, suggests that it is possible for rewarding species to evolve from saprophilous systems.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 257–268.     

Reproductive strategies of Strobilanthes kunthianus, an endemic, semelparous species in southern Western Ghats, India

Strobilanthes kunthianus (Acanthaceae) is a semelparous species with synchronized flowering and mast seeding once in 12 years. As semelparous plants have only one chance to reproduce, they are expected to develop effective strategies to prevent reproductive failure. The reproductive strategies of S. kunthianus were investigated by studying the floral traits, pollination biology, and breeding system that are critical for reproductive success. The species exhibits a series of floral traits: (1) gregarious flowering attracts a large number of Apis cerana indica , the major pollinator; (2) the stigma is sensitive to touch by the pollinator; in fresh flowers, the receptive surface faces the entry path of the incoming bee, facilitating pollen deposition; as an immediate response, the stigma curves backwards moving the receptive surface away from the path of the exiting bee, thus preventing autogamy and interference in pollen transfer; (3) flowers remain fresh for 2 days with receptive stigma and nectar and pollen reward. These traits render the species 100% pollination efficient to ensure a high seed set. As the species is self-compatible, the prevailing high degree of geitonogamous pollinations does not interfere with fruit set. The evolution of the adaptive floral traits has facilitated mast seeding in the species.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 155–163.     

Diversity, flower visitation frequency and generalism of pollinators in temperate rain forests of Chiloé Island, Chile

Species richness and taxonomic composition of pollinator assemblages are documented for 26 plant species from temperate rain forests of northern Chiloé Island, southern Chile (42°30'S). We investigated the patterns of generalism and specialization among plants and animal pollinators by comparing the flower visit frequency by different pollen vectors during the spring and summer months of three consecutive years (2000–2002). Species studied exhibited a range of floral morphologies (radial vs. zygomorphic, open vs. tubular) and rewards (nectar and/or pollen). Overall, we recorded 172 pollinator species, with an average of 6.6 species of pollen vectors/plant species. Pollinators visited an average of 15.2 plant species/pollen vector. Pollinator assemblages were dominated by Coleoptera (75 species), Diptera (56 species) and Hymenoptera (30 species), but passerine birds and hummingbirds were also important. The most specialized plants were vines, including the bee-pollinated genus Luzuriaga (Philesiaceae) and two endemic species of hummingbird-pollinated Gesneriaceae. Hymenoptera contributed 41.2% of all visits, with the bumblebee Bombus dalhbomii accounting for 22.5% of these. Plants with unspecialized flower morphology supported a higher species richness of pollinators, but visiting rates did not differ from specialized flowers.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 399–416.     

How flower biology and breeding system affect the reproductive success of the narrow endemic Polygala vayredae Costa (Polygalaceae)

Polygala vayredae is a narrow endemic species from the oriental pre-Pyrenees. Despite its conservation status and rarity, no information is available on its reproductive biology. As the flower is the structure directly involved in pollinator attraction, its morphological and functional traits have major effects on the reproductive success of the plant. In this work, the flower biology and breeding system of P. vayredae were studied to evaluate how they affect the reproductive outcome in natural populations. Flower morphology, flower rewards, and male and female functioning throughout the lifespan of the flower were assessed. Pollination experiments, involving pollinator exclusion and pollen from different sources, were conducted, and the pollen ovule index was determined. Female fitness and the occurrence of pollen limitation were assessed in three natural populations over 2 years by observing the presence of pollen on the stigma, pollen tube development, and fruit production. Polygala vayredae flowers are elaborate and long-lived with nectar rewards. The floral traits are well adapted to xenogamy and entomophily, which are in accordance with the observed breeding system and auto-incompatibility system. No mechanism of reproductive assurance was observed and P. vayredae strictly depends on pollinators to set fruit. Low fruit production was observed in the studied populations, which was largely the result of scarce, unreliable, and/or inefficient pollinators and poor pollen quality. In addition, available resources may be a limiting factor. The reproductive strategy of P. vayredae prevents inbreeding depression by a self-incompatibility system, which in years of scarce pollinators is overcome by the plant habit.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 67–81.     

Phenotypic selection in hawkmoth-pollinated Platanthera bifolia: targets and fitness surfaces

Abstract.— The present study explored phenotypic selection on phenological and morphological reproductive traits in hawkmoth-pollinated Platanthera bifolia (Orchidaceae), a Eurasian perennial herb displaying bisexual, long-spurred flowers. The work was carried out during three flowering seasons (1993–1995) in a Swedish population. Fitness was estimated as the number of pollinia removed (male fitness) and fruits produced (female fitness). Targets and patterns of selection were compared between years and sex functions by the use of multiple linear regression (including correlational selection estimates, i.e., of combination of traits), analysis of covariance, and projection pursuit regression (PPR). Results from the nonparametric surface-fitting-method PPR showed that selection was mostly linear, thus justifying the use of the parametric methods. In all study years, male and female fitness were highest in plants with many flowers. This reflects that flower number sets an upper limit to fitness and that a large inflorescence attracts more pollinators. In 1994, the summer was dry and the average spur length of P. bifolia was shorter than in the other years. In this year, male and female fitness were positively related to spur length, apparently because the spur of short-spurred plants was somewhat too short relative to the tongue length of the local pollinator for optimal pollen export and import. Additionally, the dry weather in 1994 caused a tendency for correlational selection, which was not found in the other years of study. Among small individuals (apparently more sensitive to drought than large ones), early-flowering plants had higher male and female fitness. The results show that patterns of selection may vary both between years and between sex functions in perennial hermaphroditic plants. The present study is one of the first to consider correlational selection in plants, which probably is of common occurrence and deserves to be investigated more.     

Flower-visiting bees and the breakdown of the tristylous breeding system of Eichhornia azurea (Swartz) Kunth (Pontederiaceae)

Populations of Eichhornia azurea (Swartz) Kunth (Pontederiaceae) are often composed of three floral morphs that differ in style length and anther position. Hence, E. azurea is described as tristylous. For sexual reproduction the plant requires specialized pollinators able to transfer pollen from flowers with long-, mid-, or short-level anthers to flowers with long-, mid-, or short-styles, respectively. This is because within a floral morph, self-incompatibility is the rule. Populations of E. azurea were studied from six localities in south-eastern Brazil (São Paulo and Minas Gerais). The frequencies of floral morphs, floral morphology and flower-visiting bees were recorded in each population. Ten different bee species were observed visiting plants. Four of them were considered generalist bees and not appropriate for legitimate pollination of the three floral morphs. Another three species were considered opportunists and only three species were considered to be potential pollinators. At the coastal region (Juréia), the plant population lacked the short-styled morph and the mid-styled morph and exhibited a semi-homostylous condition, in which the low-level anthers were adjacent to the stigma. This morph showed high level of self-compatibility. The disappearance of the short-styled morph from this region is likely correlated with the absence of Ancyloscelis gigas Friese, a specialized long-tongued bee that is oligolectic on this plant in other localities.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 77 , 499–507.     

Noncorrelated evolution between herbivore- and pollinator-linked features in Aristolochia chilensis (Aristolochiaceae)

The effect of mutualists (i.e. pollinators) and antagonists (i.e. herbivores) can have non-additive effects on plant fitness. This is often interpreted as evidence for correlated evolution on a suite of traits leading to an increase and decrease of the interaction of plants with mutualists and antagonists, respectively. This situation has been found to prevail in plants that have large floral and fruit displays but are not limited by pollinators for seed set. We suggest the alternative hypothesis, where plants limited by pollinators for seed set (e.g. deceit-pollinated plants) exhibit additive effects of pollinators and herbivores on fitness (i.e. noncorrelated evolution). Using a 2 × 2 factorial design, we tested this hypothesis by solely and simultaneously evaluating the effects of pollinators and the single herbivore, Battus polydamas archidamas , on female reproductive success of Aristolochia chilensis . Plants exposed to herbivores presented 2.6-fold greater herbivory than plants that excluded them. In addition, plants exposed to pollinators showed strong limitation by pollinators for seed set compared with other plants of the genus Aristolochia. However, only pollinators had a significant effect on fruit and seed set because plants that excluded them did not set fruits or seeds. Furthermore, herbivores and pollinators exerted additive effects on fruit and seed production. Collectively, these results indicate that herbivore- and pollinator-linked traits in A. chilensis exhibit noncorrelated evolution.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 239–245.     

Pollination of the lady's slipper Cypripedium henryi Rolfe (Orchidaceae)

The pollination ecology of Cypripedium henryi Rolfe, a slipper orchid endemic to west China, was investigated, and its floral shape, size, colour, and scent were analysed. Examination of the breeding system suggests that the flowers are self-compatible, but need pollen vectors for successful reproduction. The flower is rewardless; over 15 insects belonging to Araneida, Hymenoptera, Diptera, Lepidoptera, and Coleoptera were recorded as flower visitors, but most only alighted or rested on the flower. In the total 32 h of observations over 2 years, female Lasioglossum bees were found to be the most frequent visitors and the only pollinators. They showed a high visitation frequency and, surprisingly, re-visited the same flowers frequently. Cypripedium henryi probably attracts pollinators visiting the flowers through general food deception (odour components, colour, false nectar guides), as well as special structures (slippery labellum, slippery staminode). Although three Lasioglossum species visited the flowers, only L. sauterum Fan et Ebmer was found with pollen. Lasioglossum flavohirtum Ebmer was large and climbed out from the entrance. Morphologically, L. sichuanense Fan et Ebmer could be considered as a potential pollinator, but the collected specimens were found to have no pollen of C. henryi on their bodies. It was speculated that the particular floral scent of C. henryi discouraged the entrance of L. sichuanense bees. Lasioglossum sauterum was matched morphologically to the flower, but not all of the visitations resulted in effective pollinations, as some flowers of C. henryi were frequently re-visited and the pollen mass had been taken away by bees on previous visitations. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 491–499.     

Pollination by fungus gnats in four species of the genus Mitella (Saxifragaceae)

The first example of pollination by fungus gnats in the eudicots is reported. The genus Mitella (Saxifragales) is characteristically produces minute, inconspicuous, mostly dull-coloured flowers with linear, sometimes pinnately branched, petals. To understand the function of these characteristic flowers, we studied the pollination biology of four Mitella species with different floral traits and different sexual expression: dioecious M. acerina , gynodioecious M. furusei var. subramosa , and hermaphroditic M. stylosa var. makinoi and M. integripetala. Flower-bagging experiments showed that wind pollination did not occur in the dioecious and gynodioecious species. Two years of observations of flower visitors at six study sites in Japan revealed that the principal pollinators of all four Mitella were specific species of fungus gnats (Mycetophilidae), which landed on the flowers with their long spiny legs settling on the petals. Characteristically, numerous pollen grains were attached to the fungus gnats in specific locations on the body. Although, on average, 1.3–2.6 fungus gnats visited each inflorescence per day, the fruit set of both bisexual and female flowers exceeded 63%. These results suggest that fungus gnats are highly efficient pollinators of Mitella spp., and that Mitella flowers are morphologically adapted to pollination by fungus gnats.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 449–460.     

Fragrance chemotypes of Platanthera (Orchidaceae) - the result of adaptation to pollinating moths?

Floral scent of Platanthera species (Orchidaceae) in S Sweden was collected by head-space adsorption methods in the field and analysed by gas chromatography-mass spectrometry (GC-MS). Variation in scent chemistry of both Platanthera bifolia and P. chlorantha was considerable: different scent chemotypes were found among individuals as well as populations. Mainly linalool, lilac aldehydes and alcohols, geraniol, and methyl benzoate distinguished the chemotypes. Because of a high individual variation, floral scent in Platanthera is not suitable as a taxonomic tool. Scent variation can be the result of differential selection from various pollinators or pollinator groups. However, in a long-spurred race of P. bifolia the scent profile was not clearly different from that of short-spurred races in spite of their different pollinators. The two species form natural hybrids and part of the variation at the individual level may he explained by interspecific introgression. Differences found among populations in different regions may be the result of random genetic drift.     

Geographical variation in floral traits of the tree tobacco in relation to its hummingbird pollinator fauna

Nicotiana glauca , a hummingbird pollinated plant, exhibits geographical variation in several floral traits. We examined whether geographical differentiation occurred for different flower characters and if this differentiation could be explained, at least in part, by the existence or abundance of different hummingbird species in the respective pollinator assemblages. The comparison between five populations showed significant variation in six floral traits and two female fitness measures. The traits that better discriminated between populations were corolla length and corolla width. There were metric correlations between corolla length and style length in all populations studied and, in four of the five populations, both corolla length and width were also correlated. Among plants in each population, seed weight was correlated positively and significantly with style exertion, suggesting that fruit quality is dependent on the degree of cross pollination. Assemblages of hummingbirds differed between populations in species composition, visitation frequencies, and bill length. Linear regression involving bill length of the more frequent hummingbird pollinators and corolla length yielded positive and significant relationships. Thus, there appears to be an adjustment between pollinators and flowers traits that have high incidence in the among population variation.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 657–667.     

Positive effects of flower abundance and synchronous flowering on pollination success,and pollinia dispersal in rewardless Changnienia amoena (Orchidaceae)

Pollination success and pollen dispersal in natural populations depend on the spatial‐temporal variation of flower abundance. For plants that lack rewards for pollinators, pollination success is predicted to be negatively related to flower density and flowering synchrony. We investigated the relationships between pollination success and flower abundance and flowering synchrony, and estimated pollinia dispersal distance in a rewardless species, Changnienia amoena (Orchidaceae). The results obtained in the present study revealed that male pollination success was negatively influenced by population size but was positively affected by population density, whereas female pollination success was independent of both population size and density. Phenotypic analysis suggested that highly synchronous flowering was advantageous through total pollination success, which is in contrast to previous studies. These results indicate that pollination facilitation rather than competition for pollinator visits occurs in this rewardless plant. The median distance of pollinia dispersal was 11.5 m (mean distance = 17.5 m), which is comparable to that of other rewardless plants but longer than for rewarding plants. However, pollen transfer occured mainly within populations; pollen import was a rare event. Restricted gene flow by pollinia and seeds probably explains the previous population genetic reporting a high degree of genetic differentiation between populations. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 477–488.     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

Lying to Pinocchio: floral deception in an orchid pollinated by long-proboscid flies

Plants that lack floral rewards may nevertheless attract pollinators if their flowers sufficiently resemble those of rewarding plants. Flowers of the South African terrestrial orchid Disa nervosa are similar in floral dimensions and spectral reflectance to those of a sympatric nectar-producing irid ( Watsonia densiflora s.l. ). Observations showed that the orchid and Watsonia share the same pollinator, a long-proboscid tabanid fly Philoliche aethiopica . These flies visited inflorescences of both species during their foraging bouts and most (64%) observed or captured on Watsonia inflorescences carried pollinaria of the orchid on their proboscides. They probe an average of 6.3 flowers on Watsonia inflorescences, but just 1.9 flowers on the Disa inflorescences, a behaviour which would strongly promote cross-pollination in the self-compatible orchid. The orchid generally achieves high levels of pollination success, with approximately 50% of flowers receiving or exporting pollen at some sites. Pollination success was also high at one site that lacked Watsonia plants, suggesting that the orchid does not have an obligate dependence on Watsonia . Its pollination system may therefore be characterized as intermediate between generalized food deception and specific floral mimicry.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 271–278.     

Trends in floral scent chemistry in pollination syndromes: floral scent composition in hummingbird-pollinated taxa

We studied an assemblage of 17 species of bird-pollinated Ecuadorian plants (from 14 angiosperm families), including taxa pollinated by short-billed (trochiline) and sickle-billed (hermit) hummingbirds. Hummingbirds are widely supposed to ignore fragrance while visiting flowers. We collected floral headspace odours in order to test the general prediction that specialist hummingbird-pollinated flowers are scentless. In nine out of 17 of these species we failed to detect any odours using gas chromatography-mass spectrometry (GC-MS), whereas the remaining eight species produced trace levels of volatile compounds. Most of these odour compounds were of terpenoid or lipoxygenase derivation and are commonly emitted by vegetative as well as floral plant tissues. Further studies will be required to determine whether these weak odours attract alternative pollinators, repel enemies or represent vestiges of a scented ancestry.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 191–199.     

Reproductive ecology of the endemic Lecocarpus pinnatifidus (Asteraceae) in an isolated population in the Galápagos Islands

Lecocarpus pinnatifidus is an endemic member of the Asteraceae occurring on only one island in the Galápagos archipelago. The capitula are large with female ray florets and male disc florets. They are self-compatible but this study suggests fruit set is pollen limited. Visits from Xylocopa darwini and other larger insect pollinators are rare, and small insects seem to be the main pollinators. Small insects carry few pollen grains and most likely mediate self-pollinations. Self-compatibility and seed set after selfing are the most common reproductive strategy in the Galápagos Islands and L. pinnatifidus seemingly fits well into this group.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 171–180.     

Reproductive biology of the invasive exotic shrub, Rhododendron ponticum L. (Ericaceae)

The reproductive biology of an exotic species will affect its ability to become naturalized and invasive in non-native habitats. Rhododendron ponticum is an ecologically damaging exotic weed in the British Isles, which spreads predominantly by seed. I investigated how inbreeding and outcrossing affect seed production and germination in a wild population of this species in Ireland. Experimental manipulations revealed low fruit and seed set when insects were excluded from flowers, suggesting that this species has limited capability for spontaneous autogamy. Hand-pollination treatments showed that, although flowers are self-compatible (with self and same plant pollen), higher levels of seed set occur following outcrossing (xenogamy). There was no significant difference in rate of germination of seeds from inbred or outcrossed treatments. The addition of xenogamous pollen to open flowers did not increase fruit or seed set, suggesting that flowers in this population are not pollen limited: native generalist pollinators, mainly bumblebees ( Bombus spp.), are providing an adequate pollinator service. This work demonstrates that outcrossing increases seed set and pollinators are required to facilitate this. Hence, generalist native pollinators can promote invasion by exotic plants. Native pollinators can clearly play an important part in alien species invasion.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 373–381.     

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

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

Processes of isolation and introgressive interplay between Platanthera bifolia (L.) Rich and P. chlorantha (Custer) Reichb. (Orchidaceae)

NILSSON, L. A., 1983. Processes of isolation and introgressive interplay between Platanthera bifolia L. Rich, and P. chlorantha (Custer; Reichb. (Orchidaceae). Platanthera bifolia and P. chlorantha are highly specialized for pollination by crepuscular and nocturnal Lepidoptera. The plants are largely sympatric, have overlapping flowering-times, and are completely interfertile. Principal differences exist in floral fragrance and column morphology. Isolation is effected by a set of integrated morphological, ethological, and seasonal mechanisms. Odour-based constancy by moths, differences in column structure, attachment of pollinaria, spurlength, nectar availability, and flowering-peaks reduce interspecific pollen transfer. The plants form a functional species pair in relation to their pollination environment and locally share the same main pollinator species.
Hybrids, which infrequently occur in areas of sympatry, emit a blend of the species-specific fragrance compounds and have intermediate column structure. They are frequently visited by-moths but suffer a serious reduction in seed production due to imperfect interaction vis-à-vis the pollen vectors. Hybrid viscidia, in an intermediate position on the column, are touched by moths' scaly and hairy palps and get insufficient attachment, evidently causing hybrids to act solely as female recipient. First backcross individuals largely join the functional range of either species which promotes rapid introgression. The two gene-pools interplay, although always at a low level. Patterns of introgressive hybridization were exemplified.     

Pollination biology of the deceptive orchid Changnienia amoena

The pollination biology of Changnienia amoena , an endangered terrestrial orchid, was investigated at two sites in the Shennongjia Mountains, Hubei, central China. The results show that the orchid is exclusively pollinated by bumblebees. In Longmenhe, Bombus ( Diversobombus ) trifasciatus is the primary effective pollinator, whereas B.  ( Tricornibombus ) imitator is the only pollinator in Guanmenshan. These two bumblebees can be treated as a functional group because they carry pollinaria on the same position on their bodies and have similar pollinating behaviours. The morphological traits of the flower adapt precisely to the pollinators. Visitation by bumblebees is mainly in the first half of the flowering period of C. amoena . Pollinarium removal and pollinia deposition take place when bumblebees withdraw from the flower. Crossing experiments show that C. amoena is a self-compatible and outcrossing species. Fruit set in this species is low and may result from limited pollinators because 87.5% of individuals set fruits under hand-pollination, but only 6–12% of individuals set fruits under natural conditions. These results imply that it is important to conserve the pollinator community together with the plants when conservation management for this endangered species is undertaken. Artificial pollination would also be a useful measure to facilitate restoration of the small populations.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 165–175.