Caught in the act: pollination of sexually deceptive trap-flowers by fungus gnats in Pterostylis (Orchidaceae)

Background and Aims Pterostylis

is an Australasian terrestrial orchid genus of more than 400 species, most of which use a motile, touch-sensitive labellum to trap dipteran pollinators. Despite studies dating back to 1872, the mechanism of pollinator attraction has remained elusive. This study tested whether the fungus gnat-pollinated Pterostylis sanguinea secures pollination by sexual deception.

Methods

The literature was used to establish criteria for confirming sexual deception as a pollination strategy. Observations and video recordings allowed quantification of each step of the pollination process. Each floral visitor was sexed and DNA barcoding was used to evaluate the degree of pollinator specificity. Following observations that attraction to the flowers is by chemical cues, experimental dissection of flowers was used to determine the source of the sexual attractant and the effect of labellum orientation on sexual attraction. Fruit set was quantified for 19 populations to test for a relationship with plant density and population size.

Key Results

A single species of male gnat (Mycetophilidae) visited and pollinated the rewardless flowers. The gnats often showed probing copulatory behaviour on the labellum, leading to its triggering and the temporary entrapment of the gnat in the flower. Pollen deposition and removal occurred as the gnat escaped from the flower via the reproductive structures. The labellum was the sole source of the chemical attractant. Gnats always alighted on the labellum facing upwards, but when it was rotated 180 ° they attempted copulation less frequently. Pollination rate showed no relationship with orchid population size or plant density.

Conclusions

This study confirms for the first time that highly specific pollination by fungus gnats is achieved by sexual deception in Pterostylis. It is predicted that sexual deception will be widespread in the genus, although the diversity of floral forms suggests that other mechanisms may also operate.     

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

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

The effect of flower position on male and female reproductive success in a deceptively pollinated tropical orchid

In many plants, including orchids, differential fruit set along the inflorescence has been attributed to pollinator behaviour. For instance, the pollinator, moving up the inflorescence, becomes satiated with the resources and leaves before visiting the upper flowers. Consequently, the pollinators do not visit flowers as frequently higher up the inflorescence. Alternatively, flower size may vary along the inflorescence, making pollination ineffective as flowers decrease in size. I tested for the presence of differential pollination along the inflorescence in a pollinator-limited tropical epiphyte, Lepanthes rupestris Stimson, and determined the likely cause of the observed pattern. As this species has inflorescences with sequential flowering, pollinator behaviour, moving up the inflorescence as in synchronous multiflowering inflorescences, can be discounted as an explanation for differential fruit set. Fruit set is shown to be more frequent at the base of the inflorescence, but male reproductive success through pollinarium removal is basically independent of flower position. Moreover, cross-pollination by hand at variable flower positions along the inflorescence results in equal fruit set, suggesting that resources are not limiting and cannot explain the cause of differential fruit production along the inflorescence in natural populations. Furthermore, flower size is shown to diminish along the inflorescence, suggesting that the pollinator(s) may be ineffective at depositing the pollinarium in the smaller higher flowers. Consequently, pollinator behaviour and its interaction with flower size, and not resource limitation, is likely to be the main cause of differential fruit set along the inflorescence in L. rupestris .  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 405–410.     

Pollination biology of Cymbidium goeringii (Orchidaceae) in China

The genus Cymbidium comprises three subgenera with ca. 50 species. Interactions between pollinators and plants have been studied in the two subgenera Cymbidium and Cyperorchis, but only a few studies in the subgenus Jensoa have been reported. Here we report on the reproductive characteristics of C. goeringii (in sub-genus Jensoa) in three populations in the southwest of Hunan Province, China, during the winter from December 2005 to March 2006. Floral phenological and morphological features, behavior of visitors, the breeding system, and fruit sets under natural conditions were studied. The flowers of C. goeringii were strongly fragrant, but did not present any rewards to the visitors. The flowering period of C. goeringii in the studied populations lasted about 40 days, and most flowers (about 60%) opened within 30 days. Flowers opened immediately when temperature increased obviously and reached to about 16 ℃. The flowering time of the pollinated flowers, unpollinated flow-ers and the flowers with pollinarium removed were similar. Two bees of Apidae, Apis cerana cerana (honeybee) and Anthophora melanognatha, were observed visiting flowers of C. goeringii, but only the honeybees performed as pollinators. The honeybees mostly visited the orchid flowers at 10:00–17:00 on sunny days with temperature above 10 ℃. A total of only thirteen visits were observed during 20 days of observation, indicating pollinators were rare. Honeybees directly landed on the upper surface of the labellum and inserted their heads into the flowers between column and labellum, while the hind legs trod on the surface of the curved downward mid-lobe of label-lum. When a honeybee landed on the labellum of a flower, the labellum moved up and down slightly. After the honeybee entered the flower further, its head might touch the foot of the column. At this time, the body of the honeybee was parallel with the upper surface of the labellum. Then the honeybee used its front legs to scratch on the callus ridges on the upper surface of the labellum, and its hind legs hooked the edges of the side lobe of label-lum, trying to exit forcibly from the flower. During exiting process, sometimes the honeybee’s body was arched owing to tension. In this case, the surface of the scutellum came into close contact with the viscidium and then pollinaria, together with the anther cap, were removed. When this honeybee visited next flower, the pollinaria would be adhered to the stigma when it arched its body during the exiting process. Three plant species flowered synchronously with C. goeringii in the studied areas, but their flowers were different with C. goeringii in color or shape. Because C. goeringii is rewardless to pollinators, the flowers probably attract visitors by olfactory stimu-lus. Breeding system experiments showed no spontaneous autogamy and pollination success relying on pollinators in C. goeringii. Artificial pollination resulted in 90% fruit set by induced autogamy, 100% by pollinating within clone, and 100% by xenogamous pollination, respectively. These results indicate that C. goeringii is highly self-compatible and the fruit production is pollen limited. Because pollinators are essential for fruit set of C. goeringii in natural habitats, protection of wild honeybee populations or apiculture is likely a simple but effective strategy to maintain the orchid populations.     

Specialized pollination by fungus gnats in the introduced population of <Emphasis Type="Italic">Aspidistra elatior</Emphasis>

The monocot genus Aspidistra comprises rhizomatous perennials that are distributed in tropical to warm temperate regions of Asia. Little is known about the pollinators of almost all the species, probably due to the inconspicuous nature of Aspidistra flowers. Nevertheless, the unusual floral morphology suggests biotic pollination, since pollen grains are hidden under each flower’s stigma. Aspidistra elatior has been suspected to have a very peculiar pollination ecology. So far, pollination by mollusks, crustaceans, or collembolans has been suspected. However, a recent study showed that A. elatior is mainly pollinated by species of fungus gnats in Kuroshima Island, southern Japan, which is its natural habitat. Here, we investigated the pollination ecology of A. elatior in Shiga Prefecture, central Japan, which is the introduced population, to reveal whether fungus gnats are also the main pollinator in the introduced population. Our study confirmed fungus gnats pollination in the investigated pollination. Furthermore, the main pollinators (i.e., Cordyla sixi and Bradysia sp.) are the same in both Kuroshima and Shiga Prefecture. Therefore, A. elatior mainly depends on a narrow taxonomic group of fungus gnats for pollination. In contrast, we failed to document any terrestrial amphipods visiting the A. elatior flowers, in spite of a relatively high fruit set in natural conditions. This fact will refute the amphipod pollination hypothesis proposed by previous studies. We consider that A. elatior is pollinated by fungus gnats through fungal mimicry, due to its superficial similarity to mushroom fruiting bodies and strong, musky floral scent.     

Parallel floral adaptations to pollination by fungus gnats within the genus Mitella (Saxifragaceae)

The widespread pattern of parallel flower evolution as an adaptation for particular pollinator agents, known as "pollination syndromes", has long drawn attention from evolutionary biologists. Here, we report parallel evolution of saucer-shaped flowers and an associated unusual pollination system within the lineage Heucherina, a group of saxifragaceous genera. Field observations reveal that 18 of 28 plant species studied are pollinated almost exclusively by fungus gnats (Mycetophilidae). Among the 18 species with a fungus-gnat pollination system, 13 have characteristic saucer-shaped flowers and are pollinated mainly by several unspecialized mycetophilid genera with short mouthparts. We performed phylogenetic analyses using nucleotide sequences of external and internal transcribed spacers of nuclear ribosomal DNA and reconstructed ancestral floral morphologies with an establishment of the model of floral character evolution under a maximum-likelihood framework. Our analysis indicates that there is significant directionality in the evolutionary shifts of floral forms in the Heucherina. The inferred phylogeny further supports four origins of saucer-shaped flowers, which is shared among 14 species that are traditionally classified into the genus Mitella. In addition, our analysis indicates the extensive polyphyly of genus Mitella, as also suggested previously. The results suggest that the flower-visiting fungus gnats have caused convergent selection for the saucer-shaped flower repeatedly evolved within Heucherina.     

A new pollination system: brood-site pollination by flower bugs in Macaranga (Euphorbiaceae)

Background and Aims Macaranga: (Euphorbiaceae) is a large genus of dioecious trees with approx.260 species. To date, only one pollination study of the genushas reported brood-site pollination by thrips in M. hullettii.In this study, the pollination system of Macaranga tanariusis reported. Methods: The study was conducted on Okinawa and Amami Islands, Japan.Flower visitors on M. tanarius were collected and their pollenload and behaviour on the flowers examined, as well as inflorescencestructure and reward for the pollinators. Key Results: The most abundant flower visitors found on the male and femaleinflorescences were Orius atratus (Anthocoridae, Hemiptera),followed by Decomioides schneirlai (Miridae, Hemiptera). Pollenload on O. atratus from flowering pistillate inflorescenceswas detected as well as from staminate flowers. Orius atratusand D. schneirlai are likely to use the enclosed chambers formedby floral bracts as breeding sites before and during floweranthesis, and feed on nectar on the adaxial surface of flowerbracts. The extrafloral nectary has a ball-shaped structureand the contained nectar is not exposed; the hemipterans piercethe ball to suck out the nectar. Conclusions: The results indicate that the plant is pollinated by flowerbugs breeding on the inflorescences. This study may be the firstreport of pollination systems in which flower bugs are the mainpollinators. Similarity of pollination systems between M. hullettiiand M. tanarius indicates that the two brood-site pollinationsystems have the same origin. The pollinator species belongsto a predacious group, whose major prey includes thrips. Thepollination system might represent a unique example of evolutionfrom predatory flower visitors feeding on the pollinators (thrips)to the main pollinators.     

Bouncy versus idles: On the different role of pollinators in the generalist Gentiana lutea L.

Generalist flowers are visited by a broad variety of insects that function as pollinators, occasional visitors and as pollen and/or nectar robbers. Moreover, among legitimate pollinators the pollination efficiency can be different. Nectar greatly affects visitor behaviour and fidelity to a certain species, influencing plant reproductive effort. In this study we have investigated a generalist system (Gentiana lutea L.), examining the role of flower visitors and quantifying the contribution of each pollinating taxon in three natural populations. In order to verify the level of generalization, we introduce an index of Pollinator Performance (PoP), based on insect visitation rate and cross-pollen transport efficiency. Our results confirm the high degree of pollinator-generalization of the study species. Nevertheless, flower visitors show various degree of pollinating performance, mainly defined by their sedentary versus dynamic behaviour. Sedentary insects enhance geitonogamous pollen transfer, which results in reduced seed set and pollen limitation. In particular, an unusual sluggish behaviour was observed in bumblebees feeding on nectar. The hexose-rich abundant nectar offered by G. lutea flowers is remarkably rich in proline and β-alanine amino acids: this composition presumably influences feeding choice and insect dynamism, likely exerting a narcotic effect on pollinators. The consequences on plant fitness are discussed in an evolutionary perspective.     

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

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

Floral biology and histochemical analysis of Vanilla edwallii Hoehne (Orchidaceae: Vanilloideae): an orchid pollinated by Epicharis (Apidae: Centridini)

The genus Vanilla is the most diverse in Vanilloideae, with ca 90 species distributed among tropical regions. Despite their economic importance, studies on pollination of Vanilla are very scarce and data on pollinators of species endemic to Brazil are lacking. Based on fieldwork and laboratory investigations, the floral biology of V. edwallii was studied. The pollinators and pollination process were recorded at the Serra do Japi reserve, state of São Paulo, southeastern Brazil, and the presence of floral reward was also investigated. Vanilla edwallii blooms in summer. The lateral inflorescences produce up to four pale green flowers. The white labellum is united to the base of the column forming a mentum. In the studied population V. edwallii is pollinated by Epicharis (Hoplepicharis) affinis, where the males exhibit a territorial behavior, defending flowers from other possible flower visitors. The pollen is deposited on the scutellum of bees when they abandon the flower. The mentum region is dry, suggesting no nectar production. The only secretory structures are osmophores dispersed on the inner surface of the lip responsible for production of a sweet fragrance, which together with color and morphology of flowers is related to bee attraction. The labellum is rich in mucilaginous cells, while the mucilaginous substance is retained inside the cells. The histochemical analysis also detected the presence of phenolic compounds and starch concentrated mainly at the adaxial surface of the lip and around the vascular bundles.     

THE EFFECTIVENESS OF FUNGUS GNATS AS POLLINATORS

Fungus gnats (Sciaridae and Mycetophilidae) are the principal pollinators of Listera cordata (L.) R. Br. (Orchidaceae) and Scoliopus bigelovii Torr. (Liliaceae) in coastal redwood forests of northern California. Although primitive diptera have generally been regarded as relatively inefficient pollinators, fruit set for both species is high: 61–78% for L. cordata (1976–1978) and 94.3–98.5% for S. bigelovii (1978–1979). Since probability of pollination per visit is low, we attribute high fruit set to the large number of gnats present at our study sites and corresponding large number of visits to flowers. The relative frequency of geitonogamous vs. xenogamous pollen flow was estimated by emasculating flowers and subsequently comparing pollen reception or fruit set of emasculates with controls. Results for both species indicate that interplant movement of pollen is common. Thus, fungus gnats can be effective pollen vectors, both in terms of overall fruit set and potential for cross-pollination.     

Selective pollination by fungus gnats potentially functions as an alternative reproductive isolation among five Arisaema species

Background and AimsInterspecific difference in pollinators (pollinator isolation) is important for reproductive isolation in flowering plants. Species-specific pollination by fungus gnats has been discovered in several plant taxa, suggesting that they can contribute to reproductive isolation. Nevertheless, their contribution has not been studied in detail, partly because they are too small for field observations during flower visitation. To quantify their flower visitation, we used the genus Arisaema (Araceae) because the pitcher-like spathe of Arisaema can trap all floral visitors.MethodsWe evaluated floral visitor assemblage in an altitudinal gradient including five Arisaema species. We also examined interspecific differences in altitudinal distribution (geographic isolation) and flowering phenology (phenological isolation). To exclude the effect of interspecific differences in altitudinal distribution on floral visitor assemblage, we established ten experimental plots including the five Arisaema species in high- and low-altitude areas and collected floral visitors. We also collected floral visitors in three additional sites. Finally, we estimated the strength and contribution of these three reproductive barriers using a unified formula for reproductive isolation.Key ResultsEach Arisaema species selectively attracted different fungus gnats in the altitudinal gradient, experimental plots and additional sites. Altitudinal distribution and flowering phenology differed among the five Arisaema species, whereas the strength of geographic and phenological isolations were distinctly weaker than those in pollinator isolation. Nevertheless, the absolute contribution of pollinator isolation to total reproductive isolation was weaker than geographic and phenological isolations, because pollinator isolation functions after the two early-acting barriers in plant life history.ConclusionsOur results suggest that selective pollination by fungus gnats potentially contributes to reproductive isolation. Since geographic and phenological isolations can be disrupted by habitat disturbance and interannual climate change, the strong and stable pollinator isolation might compensate for the weakened early-acting barriers as an alternative reproductive isolation among the five Arisaema species.     

Plasticity in flower size as an adaptation to variation in pollinator specificity

1. Mutualisms, including plant-pollinator interactions, are an important component of ecosystems. 2. Plants can avoid the costs of variation in pollinator benefit by maintaining specificity. 3. We hypothesise a novel mechanism to ensure specificity, which takes advantage of the cognitive abilities of specific pollinators to exclude non-specific flower visitors. 4. Inflorescences of the tropical vine genus Psiguria produce flowers at regular intervals, with subsequent flowers smaller than predecessors. 5. The principle pollinators, Heliconius spp., possess an excellent spatial memory. 6. Therefore, decreasing flower size may ensure specific pollination: once Heliconius individuals have learnt the location of an inflorescence they will return, but inconspicuous flowers should reduce visits by non-specific pollinators with poorer spatial memories. 7. We tested the predictions of this hypothesis with field experiments in Panama. We confirmed that flowers on inflorescences are smaller than their predecessors. 8. Paired experiments showed that larger flowers attracted more pollinators and that the presence of an initial large flower increased subsequent visitation by Heliconius spp. to small flowers, indicating learning behaviour. 9. These results suggest that learning behaviour and decreasing flower size maintain visits from specific pollinators while reducing those from non-specific pollinators. We propose this as a novel mechanism for promoting pollinator specificity and discuss its ecological significance.     

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.     

Pollination of Oncocyclus irises (Iris: Iridaceae) by night-sheltering male bees

Irises in the section Oncocyclus (Siems.) Baker ( IRIS: Iridaceae) grow throughout the Middle East and have large and dark-coloured flowers but no nectar reward available to flower visitors. Consequently, no reward-collecting pollinators have been observed visiting the flowers during daytime. The only visitors are solitary male bees ( Eucera spp.: Apidae) that enter the flowers at dusk and stay there overnight. Here we describe the mating system of Oncocyclus irises, and the role of night-sheltering male bees in their pollination system. Pollen viability in I. haynei on Mt. Gilboa was very high (>90%) throughout all floral life stages. Stigmas were receptive in buds and in open flowers, but not in older ones. Self-pollination yielded no fruits in three species, confirming complete self-incompatibility in Oncocyclus irises. On average, 1.9 flowers were visited by each male bee before it settled for the night in the last one. Moreover, Iris pollen was present on the dorsal side of 38.8% of males caught sheltering in flower models mounted near an I. atrofusca population, indicating that pollen is transferred between flowers by night-sheltering solitary male bees. We have surveyed 13 flowering populations of six Oncocyclus species for the presence of night-sheltering male bees as well as for fruit set. We found a positive correlation, indicating that sexual reproduction in Oncocyclus irises is dependent on night-sheltering solitary male bees. Based on their complete self-incompatibility, the absence of nectar-collecting visitors during the day, and the transfer of pollen grains by the night-sheltering solitary male bees, we conclude that fertilization of Oncocyclus irises is totally dependent on pollination by night-sheltering solitary male bees.     

Intersexual mimicry and flowering phenology facilitate pollination in a dioecious habitat specialist species, <Emphasis Type="Italic">Myristica fatua</Emphasis> (Myristicaceae)

Myristica fatua is a dioecious specialist species restricted to the endangered, freshwater Myristica swamp forests in the Western Ghats, India. Earlier studies have alluded to pollination by deception in members of the Myristica genus, and thus we examined the pollination ecology comprising floral biology, flower production, flower visitors, and reproductive success in M. fatua and inferred the potential strategies that could permit such deception in this habitat specialist tree. Male flowers provide pollen rewards for an extended period of time while female flowers are rewardless and both sexes are visited by generalist insects, mainly by honeybees and stingless bees. Bee visits were significantly more frequent and longer on male than on female flowers as bees collected pollen from male flowers. We found that flower production patterns create a preponderance of males compared to females in the swamp populations. Using a model of honeybee color vision, we found the distance between the color loci of male and female flowers and based on minimum visual angle subtended by these flowers, we suggest that the two floral sexes cannot be discriminated by bees. Bees are likely deceived by the perceptual similarity of rewardless female flowers to pollen-offering male flowers and pollination is the consequence of foraging errors made by pollinators that encounter largely male–rarely female flower mosaics as they forage among clump-distributed M. fatua trees in the swamp habitat.     

A new species of Aspidistra (Ruscaceae s.l., Asparagales) from southern Vietnam, field observations on its flowering and possible pollination by flies (Phoridae)

The genus Aspidistra with almost 100 species is the most diverse in southern China and northern Vietnam. We describe a new species Aspidistra phanluongii N.Vislobokov from southern Vietnam. Species of Aspidistra are herbs characteristic of Southeast Asian forests. Flowering of these plants can be considered cryptic. There are no recorded field observations on phenology, biology of flowering or pollination of Aspidistra. Previously proposed pollinators of Aspidistra include collembolans, amphipods, fungus gnats (Diptera: Mycetophilidae, Sciaridae) and even slugs. We present observation data on the flowering of A. phanluongii in the wild. Flowers were visited by flies of Megaselia (Phoridae) and ants. Megaselia flies, well-known pollinators of numerous tropical plant species, are shown to be likely pollinators of Aspidistra phanluongii. The present study does not provide evidence of pollination by fungus gnats.     

蝶形花植物猪屎豆自花传粉机制研究

猪屎豆( Crotalaria pallida)为典型的蝶形花植物,分布极广,是路边或遭破坏生境中最常见的先锋种之一,野外观察未发现其有营养生殖的现象,主要为种子繁殖。该研究通过对自然生境中猪屎豆开花物候、访花昆虫及繁育系统的研究,旨在从繁殖的角度阐述其快速扩张的能力。结果表明：尽管猪屎豆的主要访花昆虫是蜜蜂,但蜜蜂的访花频率极低[(1.73±1.30)次/花序·h-1],且在整个花期内花药和柱头均被龙骨瓣包裹,蜜蜂访花时未成功接触柱头和花粉,不能实现传粉,因此蜜蜂不是猪屎豆有效的传粉昆虫,这与假说“蝶形花普遍是对膜翅目昆虫,尤其是对蜜蜂传粉的适应”不一致。人工授粉结果显示,猪屎豆为自交亲和种,不存在无融合生殖现象,其繁殖主要通过主动自交生产种子来实现,且在自交过程中长短花药都参与主动自交。这种自交方式不同于其他蝶形花植物的主动自交仅由短花药实现。对猪屎豆而言,长短花药均参与自交能够增加柱头的授粉几率,保证其在不利的生境中成功结籽,是其成功扩张的关键因素之一。     

Nocturnal Versus Diurnal Pollination Success in Isertia laevis (Rubiaceae): A Sphingophilous Plant Visited by Hummingbirds

Abstract: Isertia laevis (Rubiaceae) possesses flowers with traits typical for the pollination syndrome of sphingophily. Diurnal flower observation showed that nine different hummingbirds (Trochilidae) and one flower piercer (Coerebidae) were frequent visitors. Their activity on the flowers peaked in the morning hours. Very low nectar volumes were found in the morning (8.00 h) in unbagged flowers. Nectar volumes, however, reached their peaks (27 μl) at night (2.00 h) in bagged, as well as in unbagged flowers. At night few individuals of sphingids were observed. Pollination experiments showed that flowers presented to nocturnal pollinators from 18.00 h to 6.00 h had low fruit set (14 %) but high seed set (59 %). Flowers accessible from 6.00 h to 18.00 h for diurnal flower visitors showed high fruit set of 63 % but low seed set of 14 %. This suggests that pollination of individual flowers is less effective during daytime. Regarding relative reproductive success, i.e., efficiency of pollination defined as fruit set x seed set, both diurnal and nocturnal pollinators, however, are equally successful. We conclude that frequently occurring, but not very effective pollinators contribute substantially to seed production, when the expected pollinators are scarce.     

Role of honey bees (Hymenoptera: Apidae) in the pollination biology of a California native plant, Triteleia laxa (Asparagales: Themidaceae)

A central focus of pollination biology is to document the relative effectiveness of different flower visitors as pollinators. Ongoing research seeks to determine the role that introduced honey bees (Apis mellifera L.) play in the pollination of both invasive and native plants. Here we report on the importance of A. mellifera as pollinators of a California native plant, Triteleia laxa Bentham. In observation plots and transect censuses, A. mellifera overwhelmingly dominated the T. laxa flower visitor assemblage. We believe the proximity to agriculture, where A. mellifera density is higher relative to areas far from agriculture, contributes to the discrepancy between A. mellifera abundance at the two sites. Although A. mellifera were inferior flower visitors qualitatively (visited less flowers per minute), they were the most frequent interactors with flowers. Furthermore, the proportion of visits to flowers on the same plant among flower visitor species did not differ, suggesting a general mechanism by which insects forage at T. laxa flowers and that A. mellifera do not cause more deleterious geitonogamy than do native pollinators. Flower visitation rates as a function of floral display size did not differ between A. mellifera and other flower visitors. The difference in the magnitude of flower visitation (largely by A. mellifera) between sites is consistent with a difference in seed set between sites. These results suggest that non-native A. mellifera bees can play an important role in the pollination of native plant species.