A generalized pollination system in the tropics: bats, birds and Aphelandra acanthus

Background and Aims

A number of different types of flower-visiting animals coexist in any given habitat. What evolutionary and ecological factors influence the subset of these that a given plant relies on for its pollination? Addressing this question requires a mechanistic understanding of the importance of different potential pollinators in terms of visitation rate (pollinator ‘quantity’) and effectiveness at transferring pollen (pollinator ‘quality’) is required. While bat-pollinated plants typically are highly specialized to bats, there are some instances of bat-pollinated plants that use other pollinators as well. These generalized exceptions tend to occur in habitats where bat ‘quantity’ is poor due to low or fluctuating bat densities.

Methods

Aphelandra acanthus occurs in tropical cloud forests with relatively high densities of bat visitors, yet displays a mix of floral syndrome characteristics, suggesting adaptation to multiple types of pollinators. To understand its pollination system better, aspects of its floral phenology and the ‘quantity’ and ‘quality’ components of pollination by its floral visitors are studied here.

Key Results

Flowers were found to open and senesce throughout the day and night, although anther dehiscence was restricted to the late afternoon and night. Videotaping reveals that flowers are visited nocturnally by bats and moths, and diurnally by hummingbirds. Analysis of pollen deposition shows that bats regularly transfer large amounts of conspecific pollen, while hummingbirds occasionally transfer some pollen, and moths rarely do so.

Conclusions

Hummingbirds and bats were comparable in terms of pollination ‘quantity’, while bats were the most effective in terms of ‘quality’. Considering these components together, bats are responsible for approx. 70 % of A. acanthus pollination. However, bats also transferred remarkably large amounts of foreign pollen along with the conspecific grains (three of four grains were foreign). It is suggested that the negative effects of interspecific pollen transfer may decrease bat ‘quality’ for A. acanthus, and thus select for generalization on multiple pollinators instead of specialization on bats.Key words: Specialization, generalization, pollinator effectiveness, hummingbirds, floral syndrome, bat pollination, chiropterophily, ornithophily, cloud forest, heterospecific pollen transfer     

Floral scent of bat-pollinated species: West Africa vs. the New World

Floral scent of seven West African bat-pollinated tree species, belonging to six families, was collected in situ from flowering individuals using headspace adsorption. The seven species shared neither any specific compounds nor any other discernible pattern in their floral scent composition. Most of the identified compounds are common in the floral scent of species pollinated by a variety of animals. Adansonia digitata (Bombacaceae) was the only African species found to have a substantial proportion of sulphur compounds in its floral scent. This feature contrasts with the sampled New World bat-pollinated plants, which frequently contain these compounds. The floral scent of Ceiba pentandra (Bombacaceae), native to both South America and Africa, contained no sulphur substances, contradicting a previous study in the New World that identified the major floral compounds as dimethyl disulphide and dimethyl trisulphide. We suggest that the differences in the floral scent of C. pentandra , including the absence of sulphur compounds in the African variety, result from the different selective regimes exerted by the Pteropotidae bats, in Africa, and Phyllostomidae bats, in the New World, that visit their flowers.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 161–168.     

Morphological and physiological divergences within Quercus ilex support the existence of different ecotypes depending on climatic dryness

Background and Aims

According to the Grant–Stebbins model of pollinator-driven divergence, plants that disperse beyond the range of their specialized pollinator may adapt to a new pollination system. Although this model provides a compelling explanation for pollination ecotype formation, few studies have directly tested its validity in nature. Here we investigate the distribution and pollination biology of several subspecies of the shrub Erica plukenetii from the Cape Floristic Region in South Africa. We analyse these data in a phylogenetic context and combine these results with information on pollinator ranges to test whether the evolution of pollination ecotypes is consistent with the Grant–Stebbins model.

Methods and Key Results

Pollinator observations showed that the most common form of E. plukenetii with intermediate corolla length is pollinated by short-billed Orange-breasted sunbirds. Populations at the northern fringe of the distribution are characterized by long corollas, and are mainly pollinated by long-billed Malachite sunbirds. A population with short corollas in the centre of the range was mainly pollinated by insects, particularly short-tongued noctuid moths. Bird exclusion in this population did not have an effect on fruit set, while insect exclusion reduced fruit set. An analysis of floral scent across the range, using coupled gas chromatography–mass spectrometry, showed that the scent bouquets of flowers from moth-pollinated populations are characterized by a larger number of scent compounds and higher emission rates than those in bird-pollinated populations. This was also reflected in clear separation of moth- and bird-pollinated populations in a two-dimensional phenotype space based on non-metric multidimensional scaling analysis of scent data. Phylogenetic analyses of chloroplast and nuclear DNA sequences strongly supported monophyly of E. plukenetii, but not of all the subspecies. Reconstruction of ancestral character states suggests two shifts from traits associated with short-billed Orange-breasted sunbird pollination: one towards traits associated with moth pollination, and one towards traits associated with pollination by long-billed Malachite sunbirds. The latter shift coincided with the colonization of Namaqualand in which Orange-breasted sunbirds are absent.

Conclusions Erica plukenetii

is characterized by three pollination ecotypes, but only the evolutionary transition from short- to long-billed sunbird pollination can be clearly explained by the Grant–Stebbins model. Corolla length is a key character for both ecotype transitions, while floral scent emission was important for the transition from bird to moth pollination.     

Pollinators,floral morphology and scent chemistry in the southern African orchid genus Schizochilus

Floral evolution often involves suites of traits, including morphology, colour and scent, but these traits are seldom analysed together in comparative studies. We investigated the associations between floral traits and pollination systems in Schizochilus, a southern African orchid genus with small nectar-producing flowers that has not been studied previously with respect to pollination biology. Field observations indicated the presence of distinct pollination systems in the four species which occur in the Drakensberg, including pollination by muscid flies in Schizochilus angustifolius, tachinid flies in Schizochilus zeyheri, various small flies in Schizochilus bulbinella and bees and wasps in Schizochilus flexuosus. Pollination success and pollen transfer efficiency clearly differed among the four species but were not correlated with the quantity of nectar rewards. Multivariate analysis of floral morphology and floral scent chemistry based on GC-MS data revealed significant differences among species as well as populations within species. The floral scent of S. angustifolius was dominated by the benzenoid compounds benzaldehyde and phenylacetaldehyde. Samples of one population of S. bulbinella were relatively similar to S. angustifolius but samples of another population were very distinct due to the occurrence of the nitrogen-containing compounds 3-methyl-butyl aldoxime (syn/anti) and the higher amounts of aliphatic esters, alcohols and acids. In contrast, the floral scent of S. flexuosus and S. zeyheri was characterized by high relative amounts of methyl benzoate. We conclude that Schizochilus has distinct, specialized pollination systems associated with subtle but significant variation in floral morphology and scent chemistry. We also caution that sampling of several populations may be required to characterize floral scent composition at the species-level in plants.     

First record of bat-pollination in the species-rich genus Tillandsia (Bromeliaceae)

Background and Aims

Bromeliaceae is a species-rich neotropical plant family that uses a variety of pollinators, principally vertebrates. Tillandsia is the most diverse genus, and includes more than one-third of all bromeliad species. Within this genus, the majority of species rely on diurnal pollination by hummingbirds; however, the flowers of some Tillandsia species show some characteristics typical for pollination by nocturnal animals, particularly bats and moths. In this study an examination is made of the floral and reproductive biology of the epiphytic bromeliad Tillandsia macropetala in a fragment of humid montane forest in central Veracruz, Mexico.

Methods

The reproductive system of the species, duration of anthesis, production of nectar and floral scent, as well as diurnal and nocturnal floral visitors and their effectiveness in pollination were determined.

Key Results

Tillandsia macropetala is a self-compatible species that achieves a higher fruit production through outcrossing. Nectar production is restricted to the night, and only nocturnal visits result in the development of fruits. The most frequent visitor (75 % of visits) and the only pollinator of this bromeliad (in 96 % of visits) was the nectarivorous bat Anoura geoffroyi (Phyllostomidae: Glossophaginae).

Conclusions

This is the first report of chiropterophily within the genus Tillandsia. The results on the pollination biology of this bromeliad suggest an ongoing evolutionary switch from pollination by birds or moths to bats.     

Floral scents of chafer-pollinated asclepiads and a potential hybrid

Floral scent is a key functional trait for pollinator attraction to flowers, but is poorly documented in many plant lineages and pollination systems. In South African grasslands, chafer beetles (Scarabaeidae: Cetoniinae), particularly Atrichelaphinis tigrina, Cyrtothyrea marginalis and Leucoscelis spp., are common floral visitors and specialized pollination by these beetles has recently been established in several asclepiad, orchid and protea species. Chafer beetles are known to be attracted by a variety of floral volatile compounds and scent has been suggested to be an important signal in these chafer-operated pollination systems. In this study, we used dynamic headspace extraction methods and coupled gas chromatography–mass spectrometry (GC–MS) to examine the chemical composition of the floral scents of seven putatively chafer-pollinated asclepiad species in the genera Asclepias, Pachycarpus and Xysmalobium. We identified 15–57 compounds in the scents of these species, of which seven were common to all species examined. The scent profiles of each species separate into discrete clusters in two dimensional space based on non-metric multidimensional scaling (NMDS), indicating clear distinctions between species and suggesting that plants may use different combinations of volatiles to attract beetles. Two plants suspected to be intergeneric hybrids were also examined. Data on pollination systems, morphology and scent chemistry are consistent with the hypothesis that these plants are hybrids between the chafer-pollinated species Asclepias woodii and Pachycarpus concolor. The results of this study are discussed in relation to the role of chafer beetles as generalist pollinators of specialized asclepiads.     

A key role for floral scent in a wasp-pollination system in Eucomis (Hyacinthaceae)

Background and Aims

Floral scent may play a key role as a selective attractant in plants with specialized pollination systems, particularly in cases where floral morphology does not function as a filter of flower visitors. The pollination systems of two African Eucomis species (E. autumnalis and E. comosa) were investigated and a test was made of the importance of scent and visual cues as floral attractants.

Methods and Key Results

Visitor observations showed that E. autumnalis and E. comosa are visited primarily by pompilid wasps belonging to the genus Hemipepsis. These wasps carry considerably more Eucomis pollen and are more active on flowers than other visiting insects. Furthermore, experiments involving virgin flowers showed that these insects are capable of depositing pollen on the stigmas of E. autumnalis, and, in the case of E. comosa, pollen deposited during a single visit is sufficient to result in seed set. Experimental hand-pollinations showed that both species are genetically self-incompatible and thus reliant on pollinators for seed set. Choice experiments conducted in the field and laboratory with E. autumnalis demonstrated that pompilid wasps are attracted to flowers primarily by scent and not visual cues. Measurement of spectral reflectance by flower petals showed that flowers are cryptically coloured and are similar to the background vegetation. Analysis of headspace scent samples using coupled gas chromatography–mass spectrometry revealed that E. autumnalis and E. comosa scents are dominated by aromatic and monoterpene compounds. One hundred and four volatile compounds were identified in the floral scent of E. autumnalis and 83 in the floral scent of E. comosa, of which 57 were common to the scents of both species.

Conclusions

This study showed that E. autumnalis and E. comosa are specialized for pollination by pompilid wasps in the genus Hemipepsis and achieve specialization through cryptic colouring and the use of scent as a selective floral attractant.Key words: Eucomis, Pompilidae, wasp pollination, breeding system, pollination syndrome, pollinator shift, floral volatile, floral filter     

Specialized pollination in the African milkweed Xysmalobium orbiculare: a key role for floral scent in the attraction of spider-hunting wasps

Specialized pollination by prey-hunting wasps is poorly documented in rewarding plants. Furthermore, the mechanisms of achieving specialization are not clear since flowers typically produce exposed nectar and have no morphological adaptations (such as long spurs) to exclude non-pollinating visitors. We investigated the pollination of Xysmalobium orbiculare and explored the functional roles of floral scent and nectar in attracting pollinators and deterring nectar robbers. Floral visitor observations showed that this milkweed is visited almost exclusively by pompilid wasps in the genus Hemipepsis. These wasps were the only insects to carry pollinia, and a cage experiment confirmed their effectiveness in removing and inserting pollinia on flowers. Hand-pollinations showed that plants are genetically self-incompatible and thus reliant on pollinators for seed set. Palatability experiments with honeybees showed that nectar is distasteful to non-pollinating insects and is therefore likely to play a functional role in deterring nectar thieves. Choice experiments in the field showed that the wasp pollinators are attracted primarily by floral scent rather than visual cues. Analysis of spectral reflectance of flowers revealed that flowers are dull colored and are unlikely to stand out from the background vegetation. We conclude that X. orbiculare is specialized for pollination by spider-hunting wasps in the genus Hemipepsis and utilizes floral scent to selectively attract its pollinators and unpalatable nectar to deter non-pollinating visitors.     

Sulphur-containing “perfumes” attract flower-visiting bats

 We tested the attractiveness of individual scent compounds of bat-pollinated flowers to their pollinators, small flower-visiting bats of the genus Glossophaga (Phyllostomidae). Twenty compounds belonging to four different chemical substance classes were tested, both in the laboratory and in the field. In the laboratory, the bats (Glossophaga soricina) approached odour sources spontaneously and without preceding experience. Without ever receiving any reward they preferred the scent of a sulphur-containing compound, dimethyl disulphide, to several other odour components emitted by bat-pollinated flowers, and to scentless controls. In the field, at La Selva station in the tropical lowland rain forest of Costa Rica, G. commissarisi were attracted by two sulphur-containing compounds, dimethyl disulphide and 2,4-dithiapentane, to visit artificial flowers filled with sugar water. Thus, in nectarivorous bats the sense of smell obviously plays an important role in searching for and localising food sources, and even single components of the scent bouquets of bat-pollinated flowers are attractive. The preference for sulphur-containing odours seems to be innate. Accepted: 9 November 1999     

Floral volatiles play a key role in specialized ant pollination

Chemical signals emitted by plants are crucial to understand the ecology and evolution of plant–animal interactions. Scent is an important component of floral phenotype and represents a decisive communication channel between plants and floral visitors. Floral volatiles promote attraction of mutualistic pollinators and, in some cases, serve to prevent flower visitation by antagonists such as ants. Despite ant visits to flowers have been suggested to be detrimental to plant fitness, in recent years there has been a growing recognition of the positive role of ants in pollination. Nevertheless, the question of whether floral volatiles mediate mutualisms between ants and ant-pollinated plants still remains largely unexplored. Here we review the documented cases of ant pollination and investigate the chemical composition of the floral scent in the ant-pollinated plant Cytinus hypocistis. By using chemical-electrophysiological analyses and field behavioural assays, we examine the importance of olfactory cues for ants, identify compounds that stimulate antennal responses, and evaluate whether these compounds elicit behavioural responses. Our findings reveal that floral scent plays a crucial role in this mutualistic ant–flower interaction, and that only ant species that provide pollination services and not others occurring in the habitat are efficiently attracted by floral volatiles. 4-oxoisophorone, (E)-cinnamaldehyde, and (E)-cinnamyl alcohol were the most abundant compounds in Cytinus flowers, and ant antennae responded to all of them. Four ant pollinator species were significantly attracted to volatiles emitted by Cytinus inflorescences as well as to synthetic mixtures and single antennal-active compounds. The small amount of available data so far suggest that there is broad interspecific variation in floral scent composition among ant-pollinated plants, which could reflect differential responses and olfactory preferences among different ant species. Many exciting discoveries will be made as we enter into further research on chemical communication between ants and plants.     

The pollination biology of Burmeistera (Campanulaceae): specialization and syndromes

The floral traits of plants with specialized pollination systems both facilitate the primary pollinator and restrict other potential pollinators. To explore interactions between pollinators and floral traits of the genus Burmeistera, I filmed floral visitors and measured pollen deposition for 10 species in six cloud forest sites throughout northern Ecuador. Nine species were primarily bat-pollinated (84-100% of pollen transfer); another (B. rubrosepala) was exclusively hummingbird-pollinated. According to a principal components analysis of 11 floral measurements, flowers of B. rubrosepala were morphologically distinct. Floral traits of all species closely matched traditional ornithophilous and chiropterophilous pollination syndromes; flowers of B. rubrosepala were bright red, lacked odor, opened in the afternoon, and had narrow corolla apertures and flexible pedicels, which positioned them below the foliage. Flowers of the bat-pollinated species were dull-colored, emitted odor, opened in the evening, and had wide apertures and rigid pedicels, which positioned them beyond the foliage. Aperture width appeared most critical to restricting pollination; hummingbirds visited wide flowers without contacting the reproductive parts, and bats did not visit the narrow flowers of B. rubrosepala. Aperture width may impose an adaptive trade-off that favors the high degree of specialization in the genus. Other floral measurements were highly variable amongst bat-pollinated species, including stigma exsertion, calyx lobe morphology, and pedicel length. Because multiple species of Burmeistera often coexist, such morphological diversity may reduce pollen competition by encouraging pollinator fidelity and/or spatially partitioning pollinator's bodies.     

Active pollination favours sexual dimorphism in floral scent

Zoophilous flowers often transmit olfactory signals to attract pollinators. In plants with unisexual flowers, such signals are usually similar between the sexes because attraction of the same animal to both male and female flowers is essential for conspecific pollen transfer. Here, we present a remarkable example of sexual dimorphism in floral signal observed in reproductively highly specialized clades of the tribe Phyllantheae (Phyllanthaceae). These plants are pollinated by species-specific, seed-parasitic Epicephala moths (Gracillariidae) that actively collect pollen from male flowers and pollinate the female flowers in which they oviposit; by doing so, they ensure seeds for their offspring. We found that Epicephala-pollinated Phyllanthaceae plants consistently exhibit major qualitative differences in scent between male and female flowers, often involving compounds derived from different biosynthetic pathways. In a choice test, mated female Epicephala moths preferred the scent of male flowers over that of female flowers, suggesting that male floral scent elicits pollen-collecting behaviour. Epicephala pollination evolved multiple times in Phyllantheae, at least thrice accompanied by transition from sexual monomorphism to dimorphism in floral scent. This is the first example in which sexually dimorphic floral scent has evolved to signal an alternative reward provided by each sex, provoking the pollinator''s legitimate altruistic behaviour.     

Extreme divergence in floral scent among woodland star species (Lithophragma spp.) pollinated by floral parasites

Backgrounds and Aims

A current challenge in coevolutionary biology is to understand how suites of traits vary as coevolving lineages diverge. Floral scent is often a complex, variable trait that attracts a suite of generalized pollinators, but may be highly specific in plants specialized on attracting coevolved pollinating floral parasites. In this study, floral scent variation was investigated in four species of woodland stars (Lithophragma spp.) that share the same major pollinator (the moth Greya politella, a floral parasite). Three specific hypotheses were tested: (1) sharing the same specific major pollinator favours conservation of floral scent among close relatives; (2) selection favours ‘private channels’ of rare compounds particularly aimed at the specialist pollinator; or (3) selection from rare, less-specialized co-pollinators mitigates the conservation of floral scent and occurrence of private channels.

Methods

Dynamic headspace sampling and solid-phase microextraction were applied to greenhouse-grown plants from a common garden as well as to field samples from natural populations in a series of experiments aiming to disentangle the genetic and environmental basis of floral scent variation.

Key Results

Striking floral scent divergence was discovered among species. Only one of 69 compounds was shared among all four species. Scent variation was largely genetically based, because it was consistent across field and greenhouse treatments, and was not affected by visits from the pollinating floral parasite.

Conclusions

The strong divergence in floral scents among Lithophragma species contrasts with the pattern of conserved floral scent composition found in other plant genera involved in mutualisms with pollinating floral parasites. Unlike some of these other obligate pollination mutualisms, Lithophragma plants in some populations are occasionally visited by generalist pollinators from other insect taxa. This additional complexity may contribute to the diversification in floral scent found among the Lithophragma species pollinated by Greya moths.     

The missing stink: sulphur compounds can mediate a shift between fly and wasp pollination systems

The radiation of the angiosperms is often attributed to repeated evolutionary shifts between different pollinators, as this process drives diversification of floral forms and can lead to reproductive isolation. Floral scent is an important functional trait in many pollination systems but has seldom been implicated as a key mechanism in pollinator transitions. In this study, we suggest a role for sulphur compounds in mediating a shift between specialized carrion-fly and pompilid-wasp pollination systems in Eucomis (Hyacinthaceae). Flowers of closely related Eucomis species pollinated by carrion flies or pompilid wasps have very similar greenish-white flowers, but differ markedly in floral scent chemistry (determined by GC–MS analysis of headspace extracts). Comparison of the floral colours of the four Eucomis species in the visual systems of flies and wasps suggests that colour plays little role in pollinator discrimination. Nectar properties and morphology also do not differ strongly between fly- and wasp-pollinated flowers. By comparing floral scent bouquets and experimentally manipulating the scent of plants in the field, we demonstrate that shifts between wasp and fly pollination in these four congeners can depend on the production or suppression of sulphur compounds (dimethyl disulphide and dimethyl trisulphide) in the fragrance bouquet. This suggests that mutations affecting the production of particular scent compounds could precipitate shifts between pollinators, independently of floral morphology, colour or nectar properties.     

A multivariate approach to post-pollination changes in the floral scent of Platanthera bifolia (Orchidaceae)

The effect of pollination on floral scent composition and production in the moth-pollinated orchid Platanthera bifolia was studied. A significant decrease in scent production was detected both two and five days after pollination. The proportion of wilted flowers was higher in pollinated compared to control plants after seven days. A principal component analysis (PCA) showed how pollination reduced scent production. All scent compounds were affected by pollination, even though some compounds had larger impact on the overall scent reduction. A classification of samples offered a quantitative test of the changes found in individual plants after pollination. The induced decrease in floral scent production following pollination is suggested to be of adaptive value for both plants and pollinators.     

Generalized pollination,floral scent chemistry,and a possible case of hybridization in the African orchid Disa fragrans

Pollination systems in orchids tend to be specialized as a consequence of restrictive floral morphology and specific advertising signals. Here we document a notable exception: Disa fragrans subsp. fragrans, a taxon from the Drakensberg Mountains of South Africa, which is pollinated by insects belonging to at least four orders (flies, beetles, bees, and moths). Pollinaria of D. fragrans are attached to the feet of these visitors and pollination thus occurs in a rather haphazard fashion. Nevertheless, its pollination success and pollen transfer efficiency are comparable to those of its close relative, Disa sankeyi, which is pollinated by a single genus of wasps. D. fragrans has an exceptionally strong floral scent: volatile emission is 19–86 μg per inflorescence per hour, which is up to 100 fold greater than in D. sankeyi. The scent bouquet is comprised of at least 46 compounds, mostly benzenoids and phenylpropanoids, which are known to be general attractants to a wide range of insects. In contrast to D. sankeyi, the flowers of D. fragrans have a high level of spectral purity (chroma) as is typical of many generalist insect-pollinated plants. At a site where D. fragrans co-occurs with D. sankeyi we found a plant with intermediate characteristics that may be a hybrid between the two taxa. The novel case of generalist pollination in D. fragrans documented here serves as an example of how floral advertising traits might evolve during an evolutionary shift from specialized to generalized pollination.     

Is the timing of scent emission correlated with insect visitor activity and pollination in long‐spurred Satyrium species?

Plants are expected to emit floral scent when their pollinators are most active. In the case of long‐tubed flowers specialised for pollination by crepuscular or nocturnal moths, scent emissions would be expected to peak during dawn. Although this classic idea has existed for decades, it has rarely been tested quantitatively. We investigated the timing of flower visitation, pollination and floral scent emissions in six long‐spurred Satyrium species (Orchidaceae). We observed multiple evening visits by pollinaria‐bearing moths on flowers of all study species, but rarely any diurnal visits. The assemblages of moth pollinators differed among Satyrium species, even those that co‐flowered, and the lengths of moth tongues and floral nectar spurs were strongly correlated, suggesting that the available moth pollinator fauna is partitioned by floral traits. Pollinarium removal occurred more frequently during the night than during the day in four of the six species. Scent emission, however, was only significantly higher at dusk than midday in two species. Analysis of floral volatiles using gas chromatography coupled with mass spectrometry yielded 168 scent compounds, of which 112 were species‐specific. The scent blends emitted by each species occupy discrete clusters in two‐dimensional phenotype space, based on multivariate analysis. We conclude that these long‐spurred Satyrium species are ecologically specialised for moth pollination, yet the timing of their scent emission is not closely correlated with moth pollination activity. Scent composition was also more variable than expected from a group of closely related plants sharing the same pollinator functional group. These findings reveal a need for greater understanding of mechanisms of scent production and their constraints, as well as the underlying reasons for divergent scent chemistry among closely related plants.     

Floral and vegetative cues in oil-secreting and non-oil-secreting Lysimachia species

Background and Aims

Unrelated plants pollinated by the same group or guild of animals typically evolve similar floral cues due to pollinator-mediated selection. Related plant species, however, may possess similar cues either as a result of pollinator-mediated selection or as a result of sharing a common ancestor that possessed the same cues or traits. In this study, visual and olfactory floral cues in Lysimachia species exhibiting different pollination strategies were analysed and compared, and the importance of pollinators and phylogeny on the evolution of these floral cues was determined. For comparison, cues of vegetative material were examined where pollinator selection would not be expected.

Methods

Floral and vegetative scents and colours in floral oil- and non-floral oil-secreting Lysimachia species were studied by chemical and spectrophotometric analyses, respectively, compared between oil- and non-oil-secreting species, and analysed by phylogenetically controlled methods.

Key Results

Vegetative and floral scent was species specific, and variability in floral but not vegetative scent was lower in oil compared with non-oil species. Overall, oil species did not differ in their floral or vegetative scent from non-oil species. However, a correlation was found between oil secretion and six floral scent constituents specific to oil species, whereas the presence of four other floral compounds can be explained by phylogeny. Four of the five analysed oil species had bee-green flowers and the pattern of occurrence of this colour correlated with oil secretion. Non-oil species had different floral colours. The colour of leaves was similar among all species studied.

Conclusions

Evidence was found for correlated evolution between secretion of floral oils and floral but not vegetative visual and olfactory cues. The cues correlating with oil secretion were probably selected by Macropis bees, the specialized pollinators of oil-secreting Lysimachia species, and may have evolved in order to attract these bees.     

Variation in floral characters,particularly floral scent,in sapromyophilous Stemona species

Flowers or inflorescences often deploy various signals, including visual, olfactory, and gustatory cues, that can be detected by their pollinators. In many plants, these cues and their functions are poorly understood. Deciphering the interactions between floral cues and pollinators is crucial for analyzing the reproductive success of flowering plants. In this study, we examined the composition of the fetid floral scents produced by several Stemona species, including nine S. tuberosa populations from across China, using dynamic headspace adsorption, gas chromatography, and mass spectrometry techniques. We compared variations in floral phenotype, including floral longevity, nectar rewards, pollinator behavior, and flower length and color among the Stemona species. Of the 54 scent compounds identified, the major compounds include fetid dimethyl disulfide, dimethyl trisulfide, 1‐pyrroline, butyric acid, p‐cresol, isoamyl alcohol, and indole. We detected striking differentiation in floral scent at both the species and population level, and even within a population of plants with different colored flowers. Floral characteristics related to sapromyophily and deceptive pollination, including flower color mimicking livor mortis and a lack of nectar, were found in five Stemona species, indicating that Stemona is a typical sapromyophilous taxon. Species of this monocot genus might employ evolutionary tactics to exploit saprophilous flies for pollination.