The floral biology and reproductive system of <Emphasis Type="Italic">Mauritia flexuosa</Emphasis> (Arecaceae) in a restinga environment in northeastern Brazil

Studies of the floral biology of the buriti palm, Mauritia flexuosa, have presented conflicting results with respect to the mechanism of pollination, indicating either cantharophily or anemophily. To resolve this question, the floral biology of M. flexuosa was studied in a coastal restinga environment in northeastern Brazil. The reproductive system was studied experimentally, and floral visitors were collected by bagging inflorescences. In this environment, M. flexuosa, a dioecious species, has several gender-specific floral features that function to attract pollinators, especially beetles. The male flowers produce large amounts of pollen as a reward, and male and female inflorescences produce similar odors that attract pollinators to female flowers, which offer only a nectar secretion as a reward. When feeding on the female flowers, the visitors frequently come into contact with the stigmata. To increase the chances of pollination, the female flowers persist longer than the male ones, and the viability of the pollen grain is very high. A curculionid beetle species of the genus Grasidius was found to be an effective pollinator. We suspect that wind also contributes to the pollination of M. flexuosa in the study area, but in a relatively minor way.     

An assessment of pollinator visitation to Banksia spinulosa

Abstract Remote photo-monitoring of Banksia spinulosa inflorescences indicated that birds, mammals and insects were regular visitors. These included the sugar glider, brown antechinus, eastern pygmy possum, eastern spinebill, honeybee and several moth species. Eastern spine-bills were recorded at all inflorescences monitored while visitation by other animals was more sporadic. Inflorescences were visited at least once every 24 h, and visitation frequency was approximately evenly distributed during the day and night. Examination of pollen tube growth from experimental treatments indicated that pollination success was similar from both nocturnal and diurnal visitors. However, nocturnal visitors were more effective at removing pollen from newly opened flowers. The behaviour of mammals at inflorescences was such that they would transfer much more pollen to flowers than other visitors, and probably contact receptive stigmas more often. Overall, mammals were considered to be slightly more effective pollinators than the more obvious daytime visitors, eastern spinebills. Although insects visited regularly, they were thought to be less effective at pollinating flowers than vertebrate visitors. Moths carried very little pollen, and the foraging behaviour of other insects was unlikely to promote much pollination.     

Pollination in Geonoma macrostachys and Three Congeners,G. acaulis,G. gracilis,and G. interrupta

A continuous 15 month study of the floral ecology of four syntopic understorey palm species of Genoma was conducted in Amazonian Peru lowland rainforest. The spicate inflorescences of G. macrostachys, G. acaulis and G. gracilis are strictly protandrous and the plants are functionally dioecious. Data suggest that in G. macrostachys and G. acaulis pollination is based on a mimicry system, the pistillate flowers mimicking the staminate ones in colour, shape and scent. Pollen-collecting meliponine bees (Hymenoptera, Apidae, Meliponinae) and pollen-feeding syrphid flies (Diptera, Syrphidae) which visit inflorescences during both sexual stages are the pollinators of G. macrostachys. Geonoma acaulis is pollinated by small pollen-feeding weevils (Coleoptera, Curculionidae, Derelomini) that visit male and female spikes. Additionally, in G. macrostachys another pollinator type, viz. euglossine bees (Hymenoptera, Apidae, Euglossinae), which are attracted and rewarded by both types of flowers may account for long-distance pollination. The palm G. gracilis shows a very distinct pollination system. Although opportunistic insect visitors are attracted to the inflorescences of this species it seems to be mainly anemophilous because pollen becomes powdery during an thesis. The branched inflorescences of G. interrupta are also protandrous, but unlike the other species of Geonoma observed, staminate and pistillate anthesis of individual flowers are, for the most, overlapping. A broad spectrum of visitors is attracted (bees, wasps, flies, and beetles), which all may act as pollinators. Outcrossing is especially encouraged during the purely female phase at the end of the flowering cycle when there are no more staminate flowers in the inflorescence. Effects on the reproductive biology and population structure of different pollination systems and breeding system are discussed.     

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.

     

Evidence for beetle pollination in the African grassland sugarbushes (Protea: Proteaceae)

Most lineages in the African genus Protea consist of species with large unscented flowers pollinated principally by birds, and several of these lineages also show evidence of shifts to rodent pollination, associated with concealed yeasty-scented flowerheads. In this study we investigated the hypothesis that brightly coloured and fruity-scented flowerheads of four Protea species (P. caffra, P. simplex, P. dracomontana and P. welwitschii) represent a novel shift from bird to insect pollination in a grassland lineage in the genus. These species are visited by a wide range of insects, but cetoniine beetles were found to be the most important pollinators because of their abundance, size and relatively pure pollen loads. Three of the four putatively insect-pollinated Protea species have flowers presented at ground level, and experiments showed that cetoniine beetles preferred inflorescences at ground level to those artificially elevated to the height of shrubs and small trees. Relative to insects, birds were infrequent visitors to all of the study species. The nectar of all the study species contained xylose, as documented previously in bird- and rodent-pollinated Protea species, suggesting that this is a phylogenetically conserved trait. However, the very low concentration of nectar (ca. 8%), short nectar-stigma distance and the fruity scent of florets appear to be traits that are associated with specialisation for pollination by cetoniine beetles.     

Intra‐floral resource partitioning between endemic and invasive flower visitors: consequences for pollinator effectiveness

1. Sympatric flower visitor species often partition nectar and pollen and thus affect each other's foraging pattern. Consequently, their pollination service may also be influenced by the presence of other flower visiting species. Ants are solely interested in nectar and frequent flower visitors of some plant species but usually provide no pollination service. Obligate flower visitors such as bees depend on both nectar and pollen and are often more effective pollinators. 2. In Hawaii, we studied the complex interactions between flowers of the endemic tree Metrosideros polymorpha (Myrtaceae) and both, endemic and introduced flower‐visiting insects. The former main‐pollinators of M. polymorpha were birds, which, however, became rare. We evaluated the pollinator effectiveness of endemic and invasive bees and whether it is affected by the type of resource collected and the presence of ants on flowers. 3. Ants were dominant nectar‐consumers that mostly depleted the nectar of visited inflorescences. Accordingly, the visitation frequency, duration, and consequently the pollinator effectiveness of nectar‐foraging honeybees (Apis mellifera) strongly decreased on ant‐visited flowers, whereas pollen‐collecting bees remained largely unaffected by ants. Overall, endemic bees (Hylaeus spp.) were ineffective pollinators. 4. The average net effect of ants on pollination of M. polymorpha was neutral, corresponding to a similar fruit set of ant‐visited and ant‐free inflorescences. 5. Our results suggest that invasive social hymenopterans that often have negative impacts on the Hawaiian flora and fauna may occasionally provide neutral (ants) or even beneficial net effects (honeybees), especially in the absence of native birds.     

Pollination biology of the Proteaceae in Australia and southern Africa

Proteaceae are most diverse in southern Africa and Australia, especially in the south-western portions of these regions. Most genera have some species in flower at all times of the year, although generally there is a preponderance of species that flower between late winter and early summer. The one genus that is an exception to this generalization is Banksia, which either has approximately the same percentage of species in flower at various times of the year (southwestern Australia) or peaks in autumn (southeastern Australia). Within particular communities, opportunities for hybridization among congeneric species are minimized by staggered flowering times, different pollen vectors and/or various incompatibility mechanisms. Birds, mammals and arthropods have been identified as visitors to the inflorescences of many Proteaceae. The most common avian visitors to the majority of genera in Australia are honeyeaters, although lorikeets, silvereyes and approximately 40 other species sometimes may be important. Sugarbirds and sunbirds are seen most frequently at inflorescences of Protea, Leucospermum and Mimetes in southern Africa, although they rarely visit other genera. In most cases, avian visitors forage in a manner that permits the acquisition and transfer of pollen. Limited evidence supports the hypothesis that birds are selective in their choice of inflorescences, responding to morphological and/or colour changes and usually visiting those inflorescences that offer the greatest nectar rewards. Arthropods may be equally selective, although it is possible that only the larger moths, bees and beetles are important pollinators, even for those plant species that rely entirely on arthropods for pollen transfer. Mammals are pollen vectors for some Proteaceae, especially those that have geoflorous and/or cryptic inflorescences. In Australia, small marsupials may be the most important mammalian pollinators, although rodents fill this niche in at least some southern African habitats. All but two genera of Proteaceae are hermaphroditic and protandrous, the exceptions being the dioecious southern African genera Aulax and Leucadendron. For hermaphroditic species, the timing of visits by animals to inflorescences is such that they not only acquire pollen from freshly opened flowers but also brush against pollen presenters and stigmas of others that have lost self-pollen and become receptive. Birds and insects (and probably mammals) generally forage in such a way as to facilitate both outcrossing and selfing. Some species are self-compatible, although many require outcrossing if viable seed is to be formed. Regardless of which animals are the major pollen vectors, fruit set is low relative to the number of flowers available, especially in Australian habitats. Functional andromonoecy of the majority of flowers is advanced as the major cause of poor fruit set. The pollination biology and breeding systems of Australian and southern African Proteaceae resemble one another in many ways, partly because of their common ancestry, but also due to convergence. Divergence is less obvious, apart from the dichotomy between dioecious and hermaphroditic genera, and differences in the levels of seed set for Australian and African species. Future studies should concentrate on identifying the most important pollinators for various Proteaceae, the manner in which their visits are integrated with floral development and factors responsible for limiting fruit set.     

Pollination ecology of Yucca elata

The pollination biology of a population of 250 Yucca elata (Liliaceae) plants was studied in southern New Mexico. Yucca elata and the prodoxid yucca moth Tegeticula yuccasella have a mutualistic association that is essential for the successful sexual reproduction of both species. However, a wide range of other invertebrate species visit flowers during the day and at night. Our aim was to quantify the role of yucca moths and other invertebrate visitors in pollination and fruit set, using manipulative field experiments. Inflorescences were bagged during the day or night (N=12 inflorescences) to restrict flower visitors to either nocturnal or diurnal groups. Yucca moths were active exclusively nocturnally during the flowering period and thus did not visit inflorescences that were unbagged during the day. None of the 4022 flowers exposed only to diurnal visitors set fruit, whereas 4.6% of the 4974 flowers exposed only to nocturnal visitors (including yucca moths) produced mature fruit. The proportion of flowers producing fruit in the latter treatment was not significantly different from unbagged control inflorescences. In a series of experimental manipulations we also determined that: (1) flowers opened at dusk and were open for two days on average, but were only receptive to pollen on the first night of opening; (2) pollen must be pushed down the stigmatic tube to affect pollination; and (3) most plants require out-cross pollination to produce fruit. The combination of these results strongly suggests that yucca moths are the only species affecting pollination in Y. elata, and that if another species was to affect pollination, it would be a rare event.     

A preliminary study on pollination biology of Omphalogramma souliei Franch. (Primulaceae), a species endemic to China

The pollination biology of Omphalogramma souliei Franch., a species endemic to southwestern China, was investigated. Floral phenology, flower visitors, pollen/ovule ratio, attractants and rewards to the visitors were observed, measured, and recorded. Bagging experiments to exclude pollinators were carried out in the wild for two years. Our results revealed five important aspects of the reproductive biology of Omphalogramma souliei. 1) The pollen-ovule ratio was 1748±233. The breeding system was self-compatible, with facultative xenogamy. 2) The pollination syndrome is entomophily, and this species could not be pollinated by wind if the pollinators were unavailable. 3) Six insect species were observed visiting the flowers of Omphalogramma souliei in the wild, of them, three species of hymenoptera, Lasioglossum sp., Heriades parvula Cockerell and Micrapis florae Fabricius, are the principal floral visitors and effective pollinators. 4) The visual attractants to the visitors are floral color and shape, the large yellow anthers, and the rewards for visitors are pollen and nectar. 5) Cleistogamy may also occur, since the anthers of some flowers dehisced before opening of the corolla. The results of floral biology and pollination characters suggest that xenogamy predominated and autogamy played an assistant role in the evolution of reproduction and breeding system of O. souliei.     

Differences in pollinator effectiveness of birds and insects visiting Banksia menziesii (Proteaceae)

Summary The effectiveness of nectarivorous birds, introduced honey bees and staphylined beetles as pollinators of Banksia menziesii was assessed. Staphylinids removed substantial amounts of pollen but did not deposit any onto stigmata. Abundance of beetles on inflorescences was related to the mean number of florets opening per day. Honey bees collecting pollen were more likely to effect pollination than those collecting nectar which only contacted stigmata when arriving or leaving an inflorescence. Nectar-foraging birds probed between florets 10.2±0.8 (±SE) times, contacting 8–16 stigmata during each probe. Bees visited inflorescences ten times more frequently than birds although they deposited only 25% of the pollen that birds did on stigmata. Fruit set was ten times greater on inflorescences visited by birds than on inflorescences visited by bees. Bees were capable of removing as much pollen as birds but, because of direct pollen transfer to birds when florets opened during foraging, actual removal was probably much less. Selection for floret opening during nectar foraging by birds may have resulted from pollen removal by non-pollinating animals, such as staphylinids.     

Reproductive biology of Syagrus coronata (Arecaceae): sex‐biased insect visitation and the unusual case of scent emission by peduncular bracts

• Several monoecious species of palms have developed complex strategies to promote cross‐pollination, including the production of large quantities of floral resources and the emission of scents that are attractive to pollinators. Syagrus coronata constitutes an interesting model with which to understand the evolution of plant reproductive strategies in a monoecious species adapted to seasonally dry forests.
• We monitored blooming phenology over 1 year, during which we also collected and identified floral visitors and putative pollinators. We identified potential floral visitor attractants by characterizing the scent composition of inflorescences as well as of peduncular bracts, during both male and female phases, and the potential for floral thermogenesis.
• Syagrus coronata produces floral resources throughout the year. Its inflorescences are predominantly visited by a diverse assortment of small‐sized beetles, whose richness and abundance vary throughout the different phases of anthesis. We did not find evidence of floral thermogenesis. A total of 23 volatile compounds were identified in the scent emitted by the inflorescences, which did not differ between male and female phases; whereas the scent of the peduncular bracts was composed of only 4‐methyl guaiacol, which was absent in inflorescences.
• The composition of floral scent chemistry indicates that this palm has evolved strategies to be predominantly pollinated by small‐sized weevils. Our study provides rare evidence of a non‐floral scent emitting structure involved in pollinator attraction, only the second such case specifically in palms. The peculiarities of the reproductive strategy of S. coronata might play an important role in the maintenance of pollination services and pollen dispersion.

     

The pollination ecology of Actaea spicata (Ranunculaceae)

Pollination ecology of Actaea spicata L. (Ranunculaceae) was studied at localities ranging from deciduous to spruce forest. The species is self–incompatible and possibly facultatively apomictic. No anemogamy seems to occur. In deciduous forest, the plant is primarily pollinated by pollen–foraging and mate–searching Byturus ochraceus (Scriba) (Coleoptera). It flowers during a period between emergence of the beetle and onset of anthesis of the prime pollen source and only larval host plant, Geum ur–banum L. (Rosaceae). The beetles sit on the suitably long anther filaments or on top of the stigma. Floral morphology promotes pollination–performing movements by the beetle. The complex floral fragrance seems to induce close–range behaviour, i.e. alighting and/or foraging. Flies become abundant visitors only shortly before anthesis is over, and their contribution is small. Large syrphids are relatively effective as pollinators, since they grasp the stamens near the base and often touch the stigma with pollen–dusted parts of their bodies. Due to lack of its food–plant, B. ochraceus is absent at spruce forest localities and northwards. At these localities flowering is delayed until the main fly appearance, and the flowers are heavily pollinated by several syrphids and muscids. At three deciduous–forest localities a second flowering took place with new, juxtaposed inflorescences on separate stalks. The distribution of such inflorescences within the population along with the distribution and abundance of pollinators among the two generations of flowers suggest that these inflorescences occur as a response to low rate of pollination in the first anthesis.     

Pollination and fruit‐set of Grevillea robusta in western Kenya

The floral visitors of silky oak, Grevillea robusta A. Cunn. ex R.Br., their foraging behaviour and their effects on fruit‐set were studied at Malava, western Kenya. Grevillea robusta is a popular tree for farm plantings in the eastern and central African highlands. Yield of seed has been disappointingly low in some areas and a lack of appropriate pollinators has been suggested as a possible cause. Investigations involved the monitoring of visitors on active inflorescences, assessment of the rewards available to potential pollinators, and exclusion experiments to establish the effects of various visitors on fruit‐set. The flowers are visited mainly by birds and insects. The likely pollinators of G. robusta are sunbirds (Nectarinia amethystina, N. cyanolaema, N. olivacea, N. superba and N. venusta) and white‐eyes (Zosterops kikuyuensis and Z. senegalensis). Very little aggressive behaviour between birds was recorded. No nocturnal pollinators were observed. Nectar was the major floral reward for pollinators, but is likely depleted by ants and honey bees, the foraging behaviour of which confirmed them to be nectar‐robbers. These insects hardly ever touched stigmas during their visits. Eighty‐nine per cent of bird visits were in the morning (07.00–10.00 hours) when nectar volume was highest. Inflorescences bagged to exclude birds set no fruits, and unmanipulated flowers and flowers bagged with self‐pollen set no fruits, indicating a self‐incompatibility mechanism. Control cross‐pollinated flowers displayed greatly increased fruit‐set (25.1%) compared with natural open‐pollination (0.9%). All these findings confirm the importance of cross‐pollen transfer to flowers and the necessity of pollinators for fruit‐set. Effective seed production requires activity of pollinators for self‐pollen removal and cross‐pollen deposition. Seed production stands for G. robusta should be established where flowering is prolific and bird pollinators are abundant.     

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.     

Of feathers and fur: Differential pollinator roles of birds and small mammals in the grassland succulent Aloe peglerae

The pollination syndrome of the genus Aloe, with gaudy inflorescences and orange–red flowers, suggests bird pollination. However, the diversity of flowering phenologies and structures suggests that generalizations within the group are currently uninformative because few studies have addressed the role of multiple pollinator guilds, especially mammals. Aloe peglerae, endemic to montane grassland of the Magaliesberg Mountains, South Africa, is primarily bird pollinated, although small mammals are recorded nocturnal visitors. We compared the independent contributions of diurnal and nocturnal visitors, i.e. birds and small mammals respectively, to reproductive success by assigning 12 flowering A. peglerae plants to each of four pollinator selective exclusion treatments: (i) no‐visitors; (ii) nocturnal‐visitors; (iii) diurnal‐visitors; and (iv) all‐visitors (control). Birds alone contributed more to fruit set (1.2–2.6 times), average seed/fruit (1.0–1.8 times) and total seed production (1.1–2.8 times) than both the all visitors (birds and small mammals combined) and the small mammal visitors respectively. The exclusion of all visitors resulted in no fruit set, suggesting that A. peglerae requires pollinators to set seed. Germination trials over six weeks and viability testing with tetrazolium staining identified no significant differences among the three treatments that produced fruit and seed. Germination success was 90–97%, indicating that seed quality of small‐mammal and bird‐pollinated plants are similar. Aloe peglerae inflorescences were visited continuously throughout the day and night, with the Cape rock‐thrush (Monticola rupestris) and dark‐capped bulbul (Pycnonotus tricolor) being the most abundant pollinators, accounting for 68.2% and 13.7% of all visits respectively in the control treatment. Small mammals, i.e. Namaqua rock mouse (Micaelamys namaquensis) and eastern rock sengi (Elephantulus myurus), while not increasing reproductive success in the absence of birds, are suitable alternative pollinators. The presence of diurnal bird and nocturnal small mammal visitors provides a diversity of pollinators in this resilient pollination syndrome.     

The pollination of the gynomonoecious Bdallophytum oxylepis (Cytinaceae,Malvales)

Bdallophytum oxylepis is a rare and endemic species belonging to the Cytinaceae family, a root holoparasitic plant in which most resources are allocated to attracting pollinators. This species is gynomonoecious with intraindividual variation in flower size and sex. Moreover, the flowers exhibit sapromyophilous traits, as do other species of Bdallophytum. Firstly, this study aimed to determine whether all floral morphs can form seeds and be pollen donors (in the case of bisexual flowers). Secondly, as this species has floral traits hypothesized to adapt to particular types of pollen vectors (carrion flies), we also studied the pollination of B. oxylepis to confirm whether the syndromes correspond to what occurs in nature. Through pollination treatments, we determined that all floral morphs are functional. By monitoring the inflorescences, we found that pollination is specialized in the studied population. Stingless bees performed pollination, as they have a high visitation rate, frequency, and constancy, and they are unique visitors that deposit pollen on the stigmas. Thus, they appear to be effective pollinators rather than carrion flies, as predicted by the syndrome. As shown here, animal–plant interaction studies can help establish a basis for conserving rare species such as holoparasites. Moreover, knowledge about the reproductive aspects of B. oxylepis reveals essential clues about its life cycle and role in maintaining native pollinators with economic and cultural value, such as stingless bees.     

Effective pollinators of Asian sacred lotus (Nelumbo nucifera): contemporary pollinators may not reflect the historical pollination syndrome

Background and Aims

If stabilizing selection by pollinators is a prerequisite for pollinator-mediated floral evolution, spatiotemporal variation in the pollinator assemblage may confuse the plant–pollinator interaction in a given species. Here, effective pollinators in a living fossil plant Nelumbo nucifera (Nelumbonaceae) were examined to test whether beetles are major pollinators as predicted by its pollination syndrome.

Methods

Pollinators of N. nucifera were investigated in 11 wild populations and one cultivated population, and pollination experiments were conducted to examine the pollinating role of two major pollinators (bees and beetles) in three populations.

Key Results

Lotus flowers are protogynous, bowl shaped and without nectar. The fragrant flowers can be self-heating during anthesis and produce around 1 million pollen grains per flower. It was found that bees and flies were the most frequent flower visitors in wild populations, contributing on average 87·9 and 49·4 % of seed set in Mishan and Lantian, respectively. Beetles were only found in one wild population and in the cultivated population, but the pollinator exclusion experiments showed that beetles were effective pollinators of Asian sacred lotus.

Conclusions

This study indicated that in their pollinating role, beetles, probable pollinators for this thermoregulating plant, had been replaced by some generalist insects in the wild. This finding implies that contemporary pollinators may not reflect the pollination syndrome.     

Flowers and Beetles in the South American Tropics

Examples of beetle pollination in the South American tropics are given. The case histories refer to representatives of the families Cyclanthaceae, Annonaceae, and Araceae. In most cases the cantharophilous syndrome does not seem to be the result of coevolution but of a one-sided adaptation of flowers to the behavior, necessities, and sense-capacities of beetle species. It is emphasized that cantharophily may not necessarily be an indication of ancientness of a flower/insect relationship. Besides primitive angiosperms, there are derived and modern groups which, in their pollination, are specialized for beetles. Carludovica palmata (Cyclanthaceae) is pollinated by small curculionids which are attracted by odors in the early morning hours. Annona coriacea (Annonaceae) and Philodendron selloum (Araceae) are pollinated by large dynastid scarab beetles in the evening hours. In all three species thermogenesis occurs. Although belonging to widely separate systematic groups, Annona flowers and Philodendron inflorescences have a similar pollination biology, which indicates that their adaptations to beetles are the result of convergent evolutionary developments.     

Male interference with pollination efficiency in a hermaphroditic orchid

Hermaphroditism can lead to both intra‐ and intersexual conflict between male and female gender functions. However, the effect that such gender conflicts have on pollination efficiency has seldom been investigated. By artificially reducing the number of available male gametes on an individual, we quantified whether male interference with pollination efficiency occurs in the self‐compatible, moth‐pollinated orchid Satyrium longicauda. We partially emasculated S. longicauda inflorescences and compared pollination success and fecundity in these plants to intact controls. Pollen in both groups of plants was colour‐labelled so that its dispersal by pollinators could be tracked directly in the field. Intact flowers on partially emasculated inflorescences exported more pollen and received more cross‐pollen and less self‐pollen than those on intact inflorescences. Proportion of fruit set per plant was similar between the two treatments; however, fruits on partially emasculated plants had proportionally more viable seeds than those on intact controls. These results provide empirical evidence that male interference with pollination efficiency can occur in a hermaphroditic plant and that such interference can compromise fecundity. The most likely mechanism for such male interference is competition for placement on the proboscis of hawkmoth pollinators. Consequently, male competition for siring success may influence the evolution of sexual systems in hermaphroditic pollinator‐dependent plants.     

Relative importance of diurnal and nocturnal pollinators for reproduction in the early spring flowering shrub Stachyurus praecox (Stachyuraceae)

Generalized pollination systems may be favored in early spring flowering plants, as during this period pollinator activity is unpredictable. Many previous studies have concentrated on the importance of diurnal visitors in pollination, and consequently, information on the contribution of nocturnal visitors to pollination in early spring is limited. This study was conducted to evaluate the relative importance of diurnal and nocturnal pollinators in the early spring flowering dioecious shrub Stachyurus praecox (Stachyuraceae), in two temperate forests in central Japan. Visitors to the female and male flowers were observed during day and night, and their relative contributions to seed set were compared. The pollinator observations revealed that the diurnal and nocturnal insects visited both male and female flowers, and that the main flower visitors were diurnal small bees and flies as well as nocturnal settling moths. The diurnal and nocturnal flower visitors also acted as pollinators, as the pollen grains of S. praecox were attached to the insects collected from the female flowers. Pollination experiments demonstrated that the contributions of diurnal pollinators to the seed set were higher than those of the nocturnal pollinators. The results of this study indicate that S. praecox has a generalized pollination system, comprising both diurnal insects and nocturnal settling moths. Although the roles of diurnal insects are more important in the pollination of S. praecox, nocturnal settling moths may have a complementary role in early spring.