Pollinator responses to variation in floral display and flower size in dioecious Sagittaria latifolia (Alismataceae)

In animal-pollinated plants with unisexual flowers, sexual dimorphism in floral traits may be the consequence of pollinator-mediated selection. Experimental investigations of the effects of variation in flower size and floral display on pollinator visitation can provide insights into the evolution of floral dimorphism in dioecious plants. Here, we investigated pollinator responses to experimental arrays of dioecious Sagittaria latifolia in which we manipulated floral display and flower size. We also examined whether there were changes in pollinator visitation with increasing dimorphism in flower size. In S. latifolia, males have larger flowers and smaller floral displays than females. Visitation by pollinators, mainly flies and bees, was more frequent for male than for female inflorescences and increased with increasing flower size, regardless of sex. The number of insect visits per flower decreased with increasing floral display in males but remained constant in females. Greater sexual dimorphism in flower size increased visits to male inflorescences but had no influence on the number of visits to female inflorescences. These results suggest that larger flower sizes would be advantageous to both females and males, and no evidence was found that females suffer from increased flower-size dimorphism. Small daily floral displays may benefit males by allowing extended flowering periods and greater opportunities for effective pollen dispersal.     

Visitation rate of pollinators and nectar robbers to the flowers and inflorescences of Tabebuia aurea (Bignoniaceae): effects of floral display size and habitat fragmentation

Large floral displays favour pollinator attraction and the import and export of pollen. However, large floral displays also have negative effects, such as increased geitonogamy, pollen discounting and nectar/pollen robber attraction. The size of the floral display can be measured at different scales (e.g. the flower, inflorescence or entire plant) and variations in one of these scales may affect the behaviour of flower visitors in different ways. Moreover, the fragmentation of natural forests may affect flower visitation rates and flower visitor behaviour. In the present study, video recordings of the inflorescences of a tree species (Tabebuia aurea) from the tropical savannah of central Brazil were used to examine the effect of floral display size at the inflorescence and tree scales on the visitation rate of pollinators and nectar robbers to the inflorescence, the number of flowers approached per visit, the number of visits per flower of potential pollinators and nectar robbers, and the interaction of these variables with the degree of landscape disturbance. Nectar production was quantified with respect to flower age. Although large bees are responsible for most of the pollination, a great diversity of flower insects visit the inflorescences of T. aurea. Other bee and hummingbird species are highly active nectar robbers. Increases in inflorescence size increase the visitation rate of pollinators to inflorescences, whereas increases in the number of inflorescences on the tree decrease visitation rates to inflorescences and flowers. This effect has been strongly correlated with urban environments in which trees with the largest floral displays are observed. Pollinating bees (and nectar robbers) visit few flowers per inflorescence and concentrate visits to a fraction of available flowers, generating an overdispersed distribution of the number of visits per inflorescence and per flower. This behaviour reflects preferential visits to young flowers (including flower buds) with a greater nectar supply.     

Fine-scale temporal dynamics of flower visitors sheds light on insect-assemblage overlap between sexes in a dioecious Ecuadorian palm

Dioecious plants generally display sexual dimorphism in male and female floral traits, potentially attracting slightly different pollinator communities. The sharing of common floral visitors between male and female flowers and their timing of visits to both sexes is of critical importance to ensure plant's reproductive success. Palm inflorescences are visited by abundant and diverse insect communities, yet the temporal patterns of insect visits on both sexes remain poorly known. We characterized the composition of a community of flower-visiting arthropods associated with the dioecious ivory palm (Phytelephas aequatorialis, Spruce) in a pre-montane forest of Ecuador. We monitored the temporal variations in insect visits along the flowering of 12 inflorescences (eight female and four male) using interception traps recovered every 4 h. We report 59 morphospecies in the arthropod community, dominated by three beetle families: Staphylinidae, Nitidulidae, and Curculionidae. Male inflorescences were more abundantly visited than female, but visitors of the later were taxonomically more diverse. Among the 16 pollinator candidates identified, nine visited both inflorescence sexes synchronously at dusk /night whereas the others did so asynchronously during the day. Our study provides new insights into the pollination mechanism of P. aequatorialis. We found evidence of differential pollinator attraction between floral sexes, which may be explained by the sexual dimorphism of both flowers. Synchronicity in dusk/night visits of both inflorescence sexes suggests a sexual synchronization of the signal used to attract pollinators.     

Floral display size in comfrey, Symphytum officinale L. (Boraginaceae): relationships with visitation by three bumblebee species and subsequent seed set

The fecundity of insect-pollinated plants may not be linearly related to the number of flowers produced, since floral display will influence pollinator foraging patterns. We may expect more visits to plants with more flowers, but do these large plants receive more or fewer visits per flower than small plants? Do all pollinator species respond in the same way? We would also expect foragers to move less between plants when the number of flowers per plant are large, which may reduce cross-pollination compared to plants with few flowers. We examine the relationships between numbers of inflorescence per plant, bumblebee foraging behaviour and seed set in comfrey, Symphytum officinale, a self-incompatible perennial herb. Bumblebee species differed in their response to the size of floral display. More individuals of Bombus pratorum and the nectar-robbing B.?terrestris were attracted to plants with larger floral displays, but B. pascuorum exhibited no increase in recruitment according to display size. Once attracted, all bee species visited more inflorescences per plant on plants with more inflorescences. Overall the visitation rate per inflorescence and seed set per flower was independent of the number of inflorescences per plant. Variation in seed set was not explained by the numbers of bumblebees attracted or by the number of inflorescences they visited for any bee species. However, the mean seed set per flower (1.18) was far below the maximum possible (4 per flower). We suggest that in this system seed set is not limited by pollination but by other factors, possibly nutritional resources.     

Survival costs and reproductive benefits of floral display in a sexually dimorphic dioecious shrub, Leucadendron xanthoconus

The evolutionary causes of sexual dimorphism in plants have not been as widely studied as in animals and the importance of sexual selection in causing dimorphism remains controversial. Sexual selection is most obvious when it favours the evolution of a trait which enhances mating success at the expense of decreased viability. We studied the relationship between floral display (number of inflorescences), pollinator attraction and plant survival in a dioecious shrub, Leucadendron xanthoconus. Pollinator attraction, measured as the number of insect pollinators, increased linearly with floral display in males. However, males with extravagant displays had a higher probability of dying. Our data suggest that male plants are undergoing selection on floral display for increased mating success counterbalanced by selection against plants with extravagant displays. Seed set in females did not increase with floral display, except at very low inflorescence numbers. Nor was female survival correlated with floral display. Because inflorescences are terminal in the species, selection for more inflorescences in males causes increased ramification, thinner terminal branches and smaller leaves. Thus vegetative dimorphism in this species appears to be caused by selection for extravagant floral display in males, but not females. Limits to dimorphism are imposed by survival costs of elaborate display.     

Sex in advertising: dioecy alters the net benefits of attractiveness in Sagittaria latifolia (Alismataceae)

The flowers and inflorescences of animal-pollinated dioecious plants are generally small and inconspicuous in comparison with outcrossing cosexual species. The net benefits of an attractive floral display may be different for dioecious compared to cosexual populations because dioecious species experience a more severe reduction in pollen delivery when pollinators forage longer on fewer individuals. Here, we develop a model that predicts the decrease in pollen delivery in dioecious relative to cosexual populations from female-female, female-male and male-male visit sequences as the number of individuals visited varies. To evaluate the predictions of our model we conducted a common garden experiment with dioecious and monoecious (cosexual) arrays of the insect-pollinated herb Sagittaria latifolia. We find that, although increasing the advertisements of floral rewards (i.e. increasing floral display) attracts more pollinators to individuals, the probability that these pollinators subsequently deliver pollen to neighbouring plants depends on sexual system. Because the number of individual plants visited per foraging trip did not increase significantly with floral display, the relative pollination success of dioecious versus monoecious populations decreases with increased floral display. We propose that this could explain why dioecy is strongly correlated with reduced floral display among angiosperm species.     

Dioecious <Emphasis Type="Italic">Clusia nemorosa</Emphasis> achieves pollination by combining specialized and generalized floral rewards

Plants often combine multiple strategies to achieve pollen transfer. The dioecious Clusia nemorosa (Clusiaceae) produces floral resin, a specialized reward for resin-collecting bees. It also exhibits floral automimicry with female flowers mimicking male flowers in order to attract pollen-collecting insects. We observed the hourly visitation frequency and behaviour of bee visitors on two rock savanna sites in French Guiana. The major strategy in pollen transfer and the variation in visitation rates among visitor species were examined. We hypothesized that the visitation rate will vary between the floral sexes and degree of this variation will differ between the two-reward systems. We found no evidence for visits being exclusively related to resin collection, which we expected to be the principal strategy in pollen transfer. Deceit pollination appeared to have minor importance and seemed to be locally associated with the site, where demand for pollen was greater. Flower visits that probably facilitated most pollen transfer, combined resin and pollen. The pollinators involved in this system collected pollen from male flowers and resin from female flowers. The fruit set was not particularly low (44.19%) so offering different rewards by different flower genders has not constrained reproductive success in this system. Pronounced variation in visitation rates between sexes was not related to the pollination mechanisms but to the demand for the two rewards. Overall demand for pollen was greater than demand for resin. Female flowers were visited much less frequently than male flowers. Peak pollen collection occurred in the morning, i.e., as soon as the resource was available, while resin was collected throughout the day. There was a local variability in demand for the two resources because both the visitation rate and the behaviour of foraging bees varied between the two observation sites.     

Pollination ecology and reproductive success in Jack-in-the-pulpit (Arisaema triphyllum) in Québec (Canada)

Pollination ecology and reproductive success of Jack-in-the-pulpit ( Arisaema triphyllum ) were studied in two natural populations in Québec, Canada. Individual A. triphyllum plants can be of three types: male, female or bisexual. In both populations studied, the presence of bisexual inflorescences was not negligible (13%), where 'female' and 'male' bisexual plants were categorised according to the relative number of stamens and ovaries. 'Male bisexual' plants produce only pollen and 'female bisexual' plants produce only fruit. Hence, A. triphyllum is a true dioecious plant, as each plant only reproduces through either the male or the female function. 'Female bisexual' plants were equivalent to female plants in terms of visitation rate by insects, fructification rate and production of berries and seeds. Neither agamospermy in female plants nor self-pollination in 'female bisexual' plants was found, thus A. triphyllum relies on insects for cross-pollination. Despite the long flowering cycle, a low visitation rate was documented: only 20–40% of inflorescences were visited, according to gender, by a mean of 1.5 insects. In this study, Mycetophilidae represented the most generically diversified and abundant family, as well as the most efficient insect pollinator, especially the genera Docosia and Mycetophila .     

Ecological constraints on fruit initiation by Calyptrogyne ghiesbreghtiana (Arecaceae): FLORAL HERBIVORY,POLLEN AVAILABILITY,AND VISITATION BY POLLINATING BATS

Failure of a flower to initiate fruit can be attributed to an insufficiency of resources, genetic incompatibilities, or ecological constraints on pollination. Flower visitors can increase fruit initiation by pollinating flowers, or decrease fruit initiation by damaging them. If, however, resources limit fruit initiation, variation in visitation may be relatively unimportant. Visitors to Calyptrogyne ghiesbreghtiana, a rain forest understory palm, leave a record of their activity by feeding on floral tissue. I recorded variation in inflorescence visitation, floral display, and fruit initiation during 7 mo over two field seasons. On average, bats visited 50-60% of flowers; katydids and scarab beetles visited and damaged the remaining 40-50%. Four species of bat (Phyllostomidae) captured near inflorescences were found to be carrying C. ghiesbreghtiana pollen. Approximately 18% of flowers visited by bats and not damaged by insects initiated fruit, whereas insect-damaged flowers set a negligible number. Some variation in fruit initiation was explained by nightly variation in the proportion of inflorescences at each site that were potential pollen donors (male-phase inflorescences). Visitors responded to variation in floral display; katydids damaged more flowers on taller inflorescences, and in one season bats visited a greater proportion of flowers on inflorescences with many flowers. However, floral display only explained a small amount of variation in visitation. A similar amount of variation in flower visitation was explained by the degree to which inflorescences were obstructed by surrounding vegetation. I used path analysis to summarize the overall effect of floral display on visitation by both pollinators and florivores and the effect of visitation and site sex ratio on fruit initiation. Because almost all flowers were visited by bats or damaged by katydids, the amount of bat visitation was strongly negatively correlated with katydid damage. In spite of the low fruit set from bat-visited flowers, variation in bat visitation explained 27-37% of the variation in fruit initiation.     

Host-mediated volatile polymorphism in a parasitic plant influences its attractiveness to pollinators

Host-plants can mediate the interactions between herbivores and their mutualists and also between parasitic plants and their mutualists. The present study reveals how a hemiparasitic plant parasitizing three host species gives rise to three distinct hemiparasite-host neighborhoods which differ in terms of volatile composition and pollinator attractiveness. The study was performed in a population of the mistletoe Tristerix verticillatus infecting three different species of hosts occurring in sympatry within a small area, thus exposing all individuals studied to similar abiotic conditions and pollinator diversity; we assessed the effect of hosts on the hemiparasites’ visual and olfactory cues for pollinator attraction. During the study period, the hemiparasite individuals were flowering but the hosts were past their flowering stage. We collected volatile organic compounds from the hemiparasite and its hosts, measured floral display characteristics and monitored bird and insect visitors to inflorescences of T. verticillatus. We showed that: (1) floral patches did not differ in terms of floral display potentially involved in the attraction of pollinators, (2) hosts and hemiparasites on each host were discriminated as distinct chemical populations in terms of their volatile chemical profiles, (3) insect visitation rates differed between hemiparasites parasitizing different hosts, and (4) volatile compounds from the host and the hemiparasite influenced the visitation of hemiparasite flowers by insects. The study showed that a species regarded as “ornithophilic” by its floral morphology was actually mostly visited by insects that interacted with its sexual organs during their visits and carried its pollen, and that host-specific plant-volatile profiles within the T. verticillatus population were associated with differential attractiveness to pollinating insects.     

Crab spiders (Thomisidae) attract insect flower‐visitors without UV signalling

1. Crab spiders (Thomisidae) indirectly affect insect flower‐visitor and flowering plant interactions by consuming and altering the behaviour of insects. 2. Although one expects insect flower‐visitors to avoid crab spiders actively, some crab spider species are known to attract flower‐visitors. Crab spiders may use UV signalling to lure potential prey to the flowers they occupy. 3. In the present study, a field experiment was conducted to examine the effects of crab spiders occupying three prairie plant species for the insect flower‐visitor community. Pollinating insects were significantly attracted to inflorescences with crab spiders compared to inflorescences without crab spiders for two plant species, and herbivorous insects were attracted to inflorescences with crab spiders for one of these plant species. The two flowering plant species with increased pollinator visitation showed increased seed weights for plants with crab spiders, indicating crab spider presence indirectly increased pollination. 4. To test the UV signalling hypothesis, inflorescences with crab spiders of one plant species were observed under both a UV‐blocking plastic and a clear plastic control. Contrary to our prediction, flower‐visitors were not more likely to land on inflorescences under the clear plastic; the UV signalling hypothesis was not supported. Other unknown explanations underlie prey attraction to inflorescences with crab spiders.     

Bee boys and fly girls: Do pollinators prefer male or female umbels in protandrous parsnip,Trachymene incisa (Apiaceae)?

Abstract Complete dichogamy occurs when temporal separation prevents any overlap in male and female function within and among flowers of one or more inflorescences. Although dichogamy may increase outcrossing and prevent inbreeding, it also results in the presentation of inflorescences with different floral resources. Pollinators may prefer one gender over the other based on these differences, which can reduce the transfer of pollen to conspecific stigmas and reduce floral resources for effective pollinators. We investigated whether the insect visitors of Trachymene incisa (Apiaceae), an Australian herb demonstrating complete protandry at the umbel level, show a preference for male or female umbels. The male phase umbels present pollen and nectar, whereas the female phase umbels offer nectar only. Therefore, we expect pollen‐collecting insects to favour male umbels, whereas insects that forage only for nectar will favour female umbels. In natural patches that exhibited a male umbel bias, insects showed a preference for male phase umbels at Agnes Banks in 2003 and at Myall Lakes in both 2003 and 2004. By contrast, insects showed no preference for umbel phases, visiting umbels at a similar frequency to which they occurred, at Agnes Banks in 2004 and at Tomago in both 2003 and 2004. This suggests spatial and temporal variation in insect preferences for umbel phases that differ in floral rewards. In experimental arrays where the umbel gender ratio was equal, there were no significant differences between male and female umbels in terms of insect visitation during a foraging trip and mean foraging time per visit. The differing patterns of preference may be due to a differential response by insects when the umbel ratios vary, where a male bias in umbel genders leads to a preference for male umbels, whereas an equal umbel gender ratio leads to equal visitation to male and female phase umbels.     

Sexual dimorphism and sex ratios of two Australian dioecious species of alpine pineapple grass, Astelia alpina var. novaehollandiae and Astelia psychrocharis (Asteliaceae)

Sexual dimorphism and male biased sex ratios have been predicted for dioecious plants experiencing the limited conditions for growth and reproduction found in many alpine environments. To test these predictions, the reproductive ecology of two congeneric, co-occurring, dioecious, clonal, species was examined in the subalpine and alpine zones of Kosciuszko National Park, southeastern Australia. Specifically, plant size (vegetative cover of plants in quadrats), floral display (number of flowers per inflorescence, number of inflorescences per quadrat) and sex ratios (proportion of females in quadrats with flowers) were examined in ten populations of Astelia psychrocharis (Asteliaceae) and nine populations of Astelia alpina var. novae-hollandiae (Asteliaceae). Sexual dimorphism did occur, with males having more flowers per inflorescence (106% more flowers for A.alpina males and 12% more for A.psychrocharis males compared to females) and more inflorescences per quadrat than females (78% more inflorescences for A.alpina males and 46% more inflorescences for A.psychrocharis males compared to females). Plant size did not differ between male and female quadrats of either species, nor were there male biased sex ratios. However, plant size was related to flowering status in A.psychrocharis with the 65 quadrats that did not flower having lower vegetative cover than the 175 flowering quadrats indicating that there may be a minimum size/ cover required prior to flowering in this species. For A.alpina, all but two of the 185 quadrats randomly sampled flowered. There was no effect of altitude on plant size and very little effect of altitude on floral display for either species, apart from a slight increase in the number of inflorescences per quadrat with increasing altitude for A.psychrocharis, and slight decrease in number of flowers per inflorescence with increasing altitude for A.alpina females.     

神农架地区典型草本群落中的昆虫访花行为比较

对神农架地区不同海拔高度(1,000–2,600 m)的典型草本群落中的昆虫访花行为进行了初步的监测和统计。在9个样地中共观察到鳞翅目、膜翅目、双翅目11科昆虫访花, 访花者的类群在不同海拔高度的群落中有所不同, 但明显与植物群落的物种组成有关。对昆虫访花频率的监测发现, 不同物种组成的草本群落中, 所有昆虫访花的总频率和不同类群的昆虫访花频率都有差异, 正在开花的物种较多的群落中通常能吸引相对较多的访花者。同一种植物在不同的群落背景中受访频率差异显著, 如花被不甚显著的酸模(Rumex acetosa)在某些群落中未观察到访花者, 在另一些群落中却有昆虫频繁访问。而花被显著的血见愁老鹳草(Geranium henryi)在各群落中都有频繁的昆虫访问。一些植物(如广布野豌豆Vicia cracca)虽然在不同群落中都有昆虫访问, 但访花者类群却有较大差异。作者认为, 在植物群落中, 不同植物的花在吸引昆虫传粉方面存在着一定的相互作用, 在研究特定植物的虫媒传粉时应重视该植物种群所处的群落背景所起的作用。     

Neither insects nor wind: ambophily in dioecious Chamaedorea palms (Arecaceae)

Pollination of Neotropical dioecious trees is commonly related to generalist insects. Similar data for non‐tree species with separated genders are inconclusive. Recent studies on pollination of dioecious Chamaedorea palms (Arecaceae) suggest that species are either insect‐ or wind‐pollinated. However, the wide variety of inflorescence and floral attributes within the genus suggests mixed pollination mode involving entomophily and anemophily. To evaluate this hypothesis, we studied the pollination of Chamaedorea costaricana, C. macrospadix, C. pinnatifrons and C. tepejilote in two montane forests in Costa Rica. A complementary morphological analysis of floral traits was carried out to distinguish species groups within the genus according to their most probable pollination mechanism. We conducted pollinator exclusion experiments, field observations on visitors to pistillate and staminate inflorescences, and trapped airborne pollen. A cluster analysis using 18 floral traits selected for their association with wind and insect pollination syndromes was carried out using 52 Chamaedorea species. Exclusion experiments showed that both wind and insects, mostly thrips (Thysanoptera), pollinated the studied species. Thrips used staminate inflorescences as brood sites and pollinated pistillate flowers by deception. Insects caught on pistillate inflorescences transported pollen, while traps proved that pollen is wind‐borne. Our empirical findings clearly suggest that pollination of dioecious Chamaedorea palms is likely to involve both insects and wind. A cluster analysis showed that the majority of studied species have a combination of floral traits that allow for both pollination modes. Our pollination experiments and morphological analysis both suggest that while some species may be completely entomophilous or anemophilous, ambophily might be a common condition within Chamaedorea. Our results propose a higher diversity of pollination mechanisms of Neotropical dioecious species than previously suggested.     

The effects of floral display on pollinator visitation vary among populations ofPhacelia linearis (Hydrophyllaceae)

Summary Individual plants in gynodioecious populations ofPhacelia linearis (Hydrophyllaceae) vary in flower gender, flower size, and flower number. This paper reports the effects of variation in floral display on the visitation behaviour of this species' pollinators (mainly pollen-collecting solitary bees) in several natural and three experimental plant populations, and discusses the results in terms of the consequences for plant fitness. The working hypotheses were: (1) that because female plants do not produce pollen, pollen-collecting insects would visit hermaphrodite plants at a higher rate than female plants and would visit more flowers per hermaphrodite than per female; and (2) that pollinator arrival rate would increase with flower size and flower number, the two main components of visual display. These hypotheses were generally supported, but the effects of floral display on pollinator visitation varied substantially among plant populations. Hermaphrodites received significantly higher rates of pollinator arrivals and significantly higher rates of visits to flowers than did females in all experimental populations. Flower size affected arrival rate and flower visit rate positively in natural populations and in two of the three experimental populations. The flower size effect was significant only among female plants in one experimental population, and only among hermaphrodites in another. The effect of flower number on arrival rate was positive and highly significant in natural populations and in all experimental populations. In two out of three experimental populations, insects visited significantly more flowers per hermaphrodite than per female and visited more flowers on many-flowered plants than on few-flowered plants, but neither effect was detected in the third experimental population. Because seed production is not pollen-limited in this species, variation in pollinator visitation behaviour should mainly affect the male reproductive success of hermaphrodite plants. These findings suggest that pollinator-mediated natural selection for floral display inP. linearis varies in space and time.     

Pollinator response to male floral display size in two Sagittaria (Alismataceae) species

One hypothesis for the evolution of maleness in plants postulates the following: male plants appear that have re-allocated resources from female parts into a larger number of male flowers, creating a larger floral display. Pollinators respond “dramatically” to the increased display, driving the spread of males in the population (Bawa, K. S., 1980, Annual Review of Ecology and Systematics 11:15–40). To test this, we measured the total number of flowers and the size of male displays in dioecious Sagittaria latifolia and monoecious S. australis. We also measured how fast visitors arrived at “target” flowers in the displays. Then we used the Cox Model, a failure time analysis procedure, to analyze the relationship between visitor arrival and display size. We found that male display sizes were somewhat larger in the dioecious than the monoecious plants, but this was due to more compressed blooming rather than to a larger total number of flowers. The visitors to both plants were similar arrays of generalist bees. Visitors did not show the predicted “dramatic” response, however, but rather a plateauing response to larger display sizes, so our results do not support the hypothesis. Reviewing the literature, we found no reports of the “dramatic” response the hypothesis asserts. Instead, relative insensitivity to display size is the rule.     

Native birds and insects,and introduced honey bees visiting Echium wildpretii (Boraginaceae) in the Canary Islands

In this paper, we report observations of flower visitors of the endemic Echium wildpretii in Tenerife, Canary Islands. This plant inhabits the high altitudinal sub-alpine zone, which is characterized by a harsh climate, low species diversity and a short growing season. Echium wildpretii is a monocarpic perennial, producing a 2–3 m column-shaped, red-flowered, nectar-rich inflorescence. Although these floral traits have previously been suggested as being typical of ornithophilous flowers, this is the first study reporting observations of native birds (Phylloscopus collybita and Serinus canarius) in addition to insects visiting the flowers for nectar. The purposes of this study were firstly to investigate levels of visitation by native birds, native insects, and introduced honey bees. Secondly, we studied the influence of floral display (plant height and number of flowers), nearest neighbours (distance and size) and local vegetation structure on visitation rate. Finally, we discuss the evolution of ornithophily in an otherwise entomophilous plant lineage. We found that the level of bird visitation was relatively high early in the flowering season, but decreased in mid/late season, while the opposite pattern was found for introduced honey bees. For native insects, the frequency of visits was similar in early and late season. Bird visits were correlated with floral display. In the early season, visitation rates of honey bees and the two most common native bee species were correlated with size of the plant or its nearest neighbours, consistent with preference patterns for larger resource patches. Since only insects visit the flowers of other species in the Echium clade, E. wildpretii appears to have evolved from a truly insect-pollinated lineage.     

Pollinator response to female and male floral display in a monoecious species and its implications for the evolution of floral dimorphism

Pollinator-mediated selection has been hypothesized as one cause of size dimorphism between female and male flowers. Flower number, ignored in studies of floral dimorphism, may interact with flower size to affect pollinator selectivity. In the present study, we explored pollinator response, and estimated pollen receipt and removal, in experimental populations of monoecious Sagittaria trifolia, in which plants were manipulated to display three, six, nine or 12 female or male flowers per plant. In this species, female flowers are smaller but have a more compressed flowering period than males, creating larger female floral displays. Overall, pollinators preferred to visit male rather than female displays of the same size. Both first visit per foraging bout and visitation rates to female displays increased with display size. However, large male displays did not show increased attractiveness to pollinators. A predicted relationship that pollen removal, rather than pollen receipt, is limited by pollinator visitation was confirmed in the experimental populations. The results suggest that the lack of selection on large male displays may affect the evolution of floral dimorphism in this species.     

Pollination and breeding system of the enigmatic South African parasitic plant Mystropetalon thomii (Mystropetalaceae): rodents welcome,but not needed

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