Fruit set in a deceptive orchid: The effect of flowering phenology,display size,and local floral abundance

We investigated the effect of flowering time, display size, and local floral density on fruit set in Tolumnia variegata, a pollination-limited orchid that offers no reward to its pollinator(s). During 1990, natural variation in flowering time, display size, and fruit set were monitored in 508 plants at one locality in Puerto Rico. The following season, orchid floral abundance per host tree (Randia aculeata) was manipulated to investigate its effect on fruit set. Four floral abundance treatments were established (700, 500, 300, and 100), each replicated four times. Flowering time was the most important trait affecting fruit set. The proportion of plants setting at least one fruit was significantly high early and late in the season, but low during the flowering peak. Thus, strong disruptive selection differential on flowering phenology was found. Display size had little effect on fruit set. A weak, but significant disruptive selection differential on display size was found. Orchid floral abundance per host tree had a significant effect on fruit set. Early in the season, T. variegata flowers with intermediate number of conspecific flowers exhibited a greater probability of setting fruit than those in host trees with fewer or more flowers. Our results show that flowering phenology may be evolutionarily unstable, possibly a consequence of the deception pollination system. Furthermore, a deception strategy would be relatively unsuccessful in populations where plants are found in either very dense or sparse patches.     

Floral morphology and reproductive success in the orchid Epipactis helleborine: regional and local across-habitat variation

 The terrestrial orchid Epipactis helleborine is a morphologically variable species with a wide distribution in Europe. It is pollinated by social wasps, and most populations show the morphological characteristics of outcrossing species. However, local predominantly selfing subspecies and varieties have been documented from drier habitats. To document geographic variation in floral morphology, ability to produce seeds through autogamy, and reproductive success in E. helleborine, we sampled 13 populations from three geographic regions along a latitudinal gradient of c. 1000 km from northern to southern Sweden. In the southernmost region, populations in dry and mesic habitats were compared. Supplemental hand-pollination was conducted to determine whether among-population variation in fruit set could be explained by differences in the natural level of pollination, and whether any relationship between floral morphology and fruit production could be explained by interactions with pollinators. Bagging experiments showed no evidence of autogamy in any of the study populations. Number of flowers, pollinia removal and fruit set varied significantly among populations but did not differ among regions. Pollinia removal was positively correlated with population size and both pollinia removal and fruit set were lower in dry than in mesic habitats. At the level of the individual plant, the number of pollinia removed increased more rapidly with flower number than did number of fruits produced. The hand-pollination experiment indicated that the positive relationship between number of flowers and fruit production was due to a higher degree of pollen limitation in plants with few flowers than in plants with many flowers. The experiment also showed that variation in the level of pollen limitation could only partly explain variation in fruit set among populations. Received November 6, 2001; accepted April 27, 2002 Published online: December 3, 2002     

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

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

Colour similarity to rewarding model plants affects pollination in a food deceptive orchid, Orchis boryi

We studied the pollination of Orchis boryi at five different locations on the Greek mainland. Orchis boryi is food deceptive and obligatorily insect pollinated. Primary pollinators were Apis mellifera and Bombus spp., which foraged on rewarding plant species nearby and visited O. boryi in between. To analyse floral colour similarity among rewarding plants and O. boryi as perceived by bees, a model of bee colour vision was employed. For each food plant an index was calculated that described the probability of a bee foraging on it to subsequently choose an orchid flower. This choice probability correlated to colour distance according to the model of bee colour vision, indicating that bees chose the deceptive orchid more frequently if they foraged on more similarly coloured species. At different sites different plant species served as models. Bees foraging on food plants from which a high choice rate to the orchid was observed visited the orchid less often after approaching it than other bees, which is likely to reflect avoidance learning. In general, the pollination syndrome appears to be a generalized form of Batesian mimicry, in which similarity to rewarding plants determines reproductive success. As expected by negative density-dependent selection, individual fruit set and pollinia export rate correlated negatively with orchid density, but were unaffected by food plant density, orchid frequency, individual variation of labellum colour, labellum size, or mouth width of the flowers.     

Does community‐level floral abundance affect the pollination success of a rewardless orchid,Calanthe reflexa Maxim.?

Decreases in pollinator abundance may particularly constrain plants that lack floral rewards, since they are poor competitors for pollinators in the plant community. Here, we documented the pollination ecology of a rewardless orchid, Calanthe reflexa Maxim., and examined effects of forest understory degradation by deer browsing on pollination success of the species in the light of a change in the abundance of neighboring flowering plants in 2010 and 2011. Bombus species were the only pollinators at each site and the flowering phenology of C. reflexa did not overlap with that of other rewarding plants. Pollinator visit rates (assessed by time‐lapse photography), and pollinia removal rate were higher in the undegraded understory site than the degraded site in both years, while the fruit set ratio did not differ between the sites in 2011. Coverage by neighboring flowering plants was extremely low in the degraded site. Our results suggest that, although its flowering phenology and consequently lower interspecific competition of C. reflexa with rewarding plants for attracting bumblebees, neighboring flowering plants may play an important role for maintaining the visitation frequency of bumblebees of C. reflexa and contribute to its pollination success.     

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

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

Do local conspecific density and floral display size influence fruit set via pollinator visitation in Orchis militaris?

Plant density varies naturally, from isolated plants to clumped individuals, and this can influence pollinator foraging behaviour and plant reproductive success. In addition, the effect of conspecific density on reproduction may depend on the pollination system, and deceptive species differ from rewarding ones in this regard, a high density being often associated with low fruit set in deceptive plants. In our study, we aimed to determine how local conspecific density and floral display size (i.e. number of flowers per plant) affect fruit set in a deceptive orchid (Orchis militaris) through changes in pollinator visitation. We measured fruit set in a natural population and recorded pollinator abundance and foraging behaviour within plots of different O. militaris densities. Detailed data were recorded for the most abundant potential pollinators of O. militaris, i.e. solitary bees. Floral display size was negatively correlated to fruit set in medium‐density plots, but uncorrelated in low‐ and high‐density plots. Plot density had no effect on solitary bee abundance and visitation, which may be due to low pollinator abundance within the study site. The proportion of visited flowers per inflorescence was negatively influenced by floral display size, which is in line with previous studies. In addition, solitary bees spent decreasing time in successive flowers within an inflorescence, and the time spent per flower was negatively affected by ambient temperature. Our results suggest that pollinator behaviour during visitation is poorly linked to pollen deposition and reproductive success in O. militaris.     

Autonomous self‐pollination in the nectarless orchid Pogonia minor

The pollination biology of the nectarless orchid Pogonia minor was investigated in central Japan. The investigation revealed that the solitary flowers failed to attract pollinators, while high rates of fruit set were observed in the natural population. Comparable levels of fruit set were obtained in bagged, artificial self‐pollinated and artificial cross‐pollinated plants, indicating that the species is not pollinator‐limited for fruit set under natural conditions. Autonomous self‐pollination in P. minor resulted from a reduced rostellum, which allowed contact between the pollinia and the stigma. Self‐pollination is thought to be an adaptive response that provides reproductive assurance under conditions of pollinator limitation. Since pollen limitation is generally known to be frequent among deceptive orchids, strong pollen limitation is probably a driving force in the autonomous self‐pollination mechanism in the nectarless orchid P. minor.     

Female and male fitness of a sexually deceptive orchid with a narrow distribution area: from phenotypic traits to spatial distribution patterns

• The Orchidaceae family presents one of the most extravagant pollination mechanisms: deception. While many studies on reproductive success have been performed on food‐deception orchids, less have been performed on sexually deceptive orchids. Here, we focused on Ophrys balearica P. Delforge, an endemic orchid of the Balearic Islands, to study its reproductive ecology, the spatio‐temporal variation of its reproductive success and the individual (floral display and geospatial position) and population parameters (patch size, shape and density) that affect its reproductive success.
• We performed hand‐pollination experiments, along with the recording of floral display parameters and GPS position of over 1,100 individuals from seven populations in two consecutive years. We applied, for the first time, GIS tools to analyse the effects of individual’s position within the population on the reproductive success. Reproductive success was measured both in male (removed pollinia) and female (fruit set) fitness.
• The results confirm that this species is pollinator‐dependent and mostly allogamous, but also self‐compatible. This species showed high values for the cumulative inbreeding depression index and high pollen limitation. Male fitness was almost equal to female fitness between years and populations, and reproductive success exhibited huge spatio‐temporal variation.
• Although we did not find strong correlations between floral display and reproductive success, patches with low‐plant density and individuals in the external portion of the population showed significantly higher plant fitness. These findings must be considered in conservation actions for endangered orchid species, especially considering that most orchids are strongly dependent on pollinators for their species’ fitness.

     

Reproductive success of Dactylorhiza incarnata ssp. incarnata (Orchidaceae): the effects of population size and plant visibility

Reproduction of plants pollinated solely by flower-visiting animals depends on the ability of the population and each of its flowering member to attract pollinators. Factors affecting the pollination of nectarless species differ somewhat from those affecting the pollination of rewarding species due to the avoidance behaviour of pollinators after visiting empty flowers.
We studied a non-mimic food-deceptive orchid, Dactylorhiza incarnata ssp. incarnata, in 16 populations in central Finland to examine if population properties and plant size affected reproductive success of plants.
We found that the number of flowering plants increased total pollinia removal and seed production of the population, but had no effect on reproduction of individual plants. Dactylorhiza incarnata occurred in open mires with virtually no rewarding species in the neighbourhood, but the distance from the forest edge did not have any effect on reproduction. However, increased variation in the number of flowers among within-population plants enhanced plant reproductive success. The increased variation probably hampered the learning process of flower visitors. Plant size affected reproductive success only in populations with a high general reproductive success indicating that in the rarely visited populations, pollination of individual plants is mainly affected by random effects.     

Mast fruiting in a hawkmoth-pollinated orchid Habenaria glaucifolia: an 8-year survey

Aims The hypothesis of predator satiation has been proposed to explain mast fruiting in various flowering plants. It considers that the simultaneous production of large numbers of seeds by a plant population reduces the risk of seed predation for each individual. Orchids produce huge numbers of seeds per fruit and rarely experience seed predation. It remains unclear which factors may affect fluctuating fruit production in orchids, which generally suffer a widespread pollen limitation. To explore the temporal pattern of fruiting and potential factors related to fluctuation in fruit production, we investigated reproductive success of a long-spurred orchid (Habenaria glaucifolia) in an alpine meadow with thousands of individuals over 8 years.Methods To estimate reproductive success, pollinator observation was conducted by day and at night, and pollinia removal and receipt were recorded in the field population for 8 years. To examine whether fruit set and seed set are pollen limited, we conducted supplementary pollination experiments and compared fruit set, seed set and pollinia movement of open-pollinated flowers from 2011 to 2013. We measured lengths of spurs and pollinator proboscises, and nectar volume and concentration, to identify potential pollinators.Important findings Hawkmoths were seen to be effective pollinators for H. glaucifolia in 3 years, whereas in the remaining 5 years no pollinators were observed, and consequently pollinia were rarely transferred. Numerous pollinia movements were observed in 2012, 2013 and 2014 (pollinia removal: 48, 59 and 85%; pollinia receipt 51, 70 and 80%), and correspondingly fruit set was significantly higher in 2012 and 2013 (59 and 46%) than in 2011 (25%). It was fruit set, rather than seed set, that was pollen limited in this orchid in the 3 years, in that supplementary pollination increased fruit set but did not increase seed set per fruit compared to natural. Three species of hawkmoths had proboscis lengths that matched the spur length of H. glaucifolia. Fruit set in this long spurred orchid depends on the activity of long-tongued hawkmoths, resulting in significant temporal variation in fruit production. Mast fruiting in this alpine orchid could be attributed to a 'sit and wait' strategy, awaiting an abundance of effective pollinators.     

Floral display size and spatial distribution of potential mates affect pollen deposition and female reproductive success in distylous Pulmonaria officinalis (Boraginaceae)

In animal‐pollinated plants, both the spatial distribution of flowering individuals and the number of flowers that an individual displays affect pollen deposition rates and female reproductive success. Heterostylous species are likely to be particularly sensitive to the contingencies of spatial distribution, as they are reproductively subdivided into distinct mating groups, which usually exhibit self‐ and intra‐morph incompatibility and differ in floral morphology. In this paper, we explore the joint effects of both spatial distribution of potential mates and floral display size on morph‐specific pollen deposition rates and seed set patterns in two natural populations of Pulmonaria officinalis, a distylous species with a weak self‐incompatibility system. Both total stigmatic pollen load and the proportion of legitimate pollen decreased with increasing spatial isolation. Legitimate (intermorph) pollen transfer was, however, asymmetric and decreased more rapidly with decreasing proximity to a compatible legitimate mating partner in the S‐morph than in the L‐morph. Total stigmatic pollen loads per flower increased with increasing floral display size, indicating that large plants are disproportionately more visited than smaller individuals. However, because legitimate pollen deposition decreased with increasing floral display size, these results also suggest that larger numbers of flowers increase the degree of geitonogamous pollination. In both the L‐ and S‐morph, seed set significantly decreased with increasing isolation from a legitimate mating partner, but in the L‐morph seed set was less dependent on the spatial distribution of the S‐morph. In addition, seed set significantly increased with floral display size in the L‐morph, but not in the S‐morph. These findings indicate that the spatial distribution of potential mates and variation in floral display size may cause morph‐specific differences in pollen deposition rates and female reproductive success.     

Effects of distance from models on the fitness of floral mimics

• Rewardless plants can attract pollinators by mimicking floral traits of rewarding heterospecific plants. This should result in the pollination success of floral mimics being dependent on the relative abundance of their models, as pollinator abundance and conditioning on model signals should be higher in the vicinity of the models. However, the attraction of pollinators to signals of the models may be partially innate, such that spatial isolation of mimics from model species may not strongly affect pollination success of mimics.
• We tested whether pollination rates and fruit set of the rewardless orchid Disa pulchra were influenced by proximity and abundance of its rewarding model species, Watsonia lepida.
• Pollination success of the orchid increased with proximity to the model species, while fruit set of the orchid increased with local abundance of the model species. Orchids that were experimentally translocated outside the model population experienced reduced pollinaria removal and increased pollinator‐mediated self‐pollination.
• These results confirm predictions that the pollination success of floral mimics should be dependent on the proximity and abundance of model taxa, and thus highlight the importance of ecological facilitation among species involved in mimicry systems.

     

Heterostyly promotes compatible pollination in buckwheats: Comparisons of intraflower,intraplant, and interplant pollen flow in distylous and homostylous Fagopyrum

Premise of the Study

Heterostyly, the reciprocal positioning of stigmas and anthers in different floral morphs, has long been thought to promote intermorph pollination. However, extensive intramorph pollination occurs commonly in heterostylous species, leading to recurrent questions about the functional and evolutionary significance of heterostyly.

Methods

To identify the sources of stigmatic pollen (autogamous [intraflower], geitonogamous [intraplant], vs. interplant), we emasculated either one flower or entire plants in experimental populations of the two closely related buckwheat species, distylous Fagopyrum esculentum and homostylous F. tataricum. Differences in pollen size allowed unambiguous identification of pollen on stigmas.

Results

Only 2.4% of F. tataricum pollen and 1.5% of F. esculentum pollen arrived successfully on compatible stigmas of other plants. In the former (homostylous) species, 71.3% of the pollen load on stigmas was autogamous, 10.8% was geitonogamous, and 17.9% was interplant. In the latter (distylous) species, 37.45% of the pollen on stigmas was autogamous, 13.8% was geitonogamous, 17.0% was intramorph, and 31.75% was intermorph. The amount of incompatible pollen arriving on stigmas was greatly decreased by both one‐flower and whole‐plant emasculations, and thus, the proportion of compatible pollen deposited increased with one‐flower emasculation and increased even more with whole‐plant emasculation.

Conclusions

Our quantification of pollen‐donor sources in these two species indicated that heterostyly in Fagopyrum esculentum provided a nearly 2‐fold fitness advantage (in terms of compatible pollination) over expected (random) pollen transfers between morphs. Because of reduced herkogamy, the homostylous F. tataricum was highly autogamous.     

Effects of conspecific and heterospecific floral density on the pollination of two related rewarding orchids

Variation in within-population floral density can affect interactions between plants and pollinators, resulting in variable pollen export for plants. We investigated the effects of conspecific and heterospecific floral densities on pollination success both of two related, self-compatible, nectar-rewarding orchid species in Ireland, Spiranthes romanzoffiana (rare and listed as endangered) and its congener, S. spiralis (more abundant and not of conservation concern). Floral densities, insect visitation rates, and orchid pollen transport were recorded in multiple quadrats in four populations of both orchid species over their flowering season. We found that conspecific and heterospecific co-flowering plant density affected pollination in both orchid species. For S. romanzoffiana, higher heterospecific density increased pollen removal. For S. spiralis, higher conspecific visitation increased pollen removal and increased heterospecific density decreased pollen deposition. In addition, increased conspecific density increased pollen deposition in both species. This study shows that plants may interact to facilitate or compete for different components of the pollination process, namely; pollinator attraction, pollen removal and deposition. Such interactions have immediate consequences for endangered plant species, as increases in both conspecific and heterospecific coflowering density may ameliorate the negative effects of rarity on pollination, hence overall reproductive success.     

What We Think We Know vs. What We Need to Know About Orchid Pollination and Conservation: <Emphasis Type="Italic">Cypripedium</Emphasis> L. as a Model Lineage

While Darwin (1862, 1877) showed that reproductive success in orchid populations depended on adaptive floral morphology coupled with pollinator visitation a more recent review of the literature (Tremblay et al., 2005) confirmed that many out-breeding species are pollinator-limited because most orchid species showing low fecundity also lack rewards. The absence of rewards depresses both pollinator fidelity and the frequency of pollinator visits to an orchid population even though orchid flowers that lack rewards retain the same interlocking floral structures for precise pollinia removal and deposition found in related species that offer rewards. Using the genus, Cypripedium, as a model lineage of non-rewarding flowers this study also shows that the correlation between low fruit set in a Cypripedium sp. and its specific pollinator(s) is insufficient to predict specific frequencies of low fecundity. Annual rates of fruit set often vary broadly between populations of the same species and within the same population over several seasons. We speculate that fruit-set rates also decline when orchid demography and additional biotic and abiotic factors interrupt rates of pollinator activity (pre-zygotic) and fertilization/fruit maturation (post-zygotic). We suggest that that traditional field studies on pollination ecology and breeding systems be combined with data sets recording genetic variation and orchid flower demography in relation to seasonal variation in climate. We also propose that the same information be collected in regard to genetic variation, demography and phenology of populations of known orchid pollinators and co-blooming angiosperm species native to orchid habitats.     

Bird pollination in an angraecoid orchid on Reunion Island (Mascarene Archipelago, Indian Ocean)

BACKGROUND AND AIMS: Although numerous angraecoid orchids in Madagascar display typical sphingophilous syndrome (i.e. white, nectariferous, long-spurred flowers, producing a strong scent at the crepuscule that is attractive to moths), three species of Angraecum in Reunion, belonging to the endemic section Hadrangis, have atypical unscented and short-spurred flowers. The aim of the study was to investigate the implication of plant-pollinator interaction on the evolution of floral morphology of these peculiar island floral forms. METHODS: The flower morphology of A. striatum (one of the three section Hadrangis species) was investigated by performing a set of floral measures, and the reproductive biology was investigated by a set of hand pollination experiments. Natural pollinators were observed by means of a digital video camera. Pollinator efficiency (pollen removal and deposition) and reproductive success (fruit set) were quantified once a week in natural field conditions during the 2005 flowering season (i.e. from January to March). KEY RESULTS: The orchid is self-compatible but requires a pollinator to achieve fruit set. Only one pollinator was observed, the endemic white-eye Zosterops borbonicus (Zosteropidae). These birds perched on inflorescences, and probed most fresh-looking flowers on each plant for nectar. Nectar was both abundant (averaging 7.7 microL) and dilute (averaging 9.7 % sugar in sucrose equivalents). Birds were mostly active between 0830 and 0930 h. Visits to plants were extremely short, lasting from 9 to 27 s. At the study site, 60.9 % of flowers had pollen removed, and 46.4 % had pollinia deposited on stigmas. The proportion of flowers that initiated a fruit averaged 20.6 % in natural conditions. CONCLUSIONS: For the first time, a bird-pollinated orchid is described from a sub-tribe that is mainly specialized for moth pollination. This study documents a morphological shift in flowers in response to pollinator adaptations in the insular context of the Mascarene Archipelago.     

Pollination biology and breeding systems in the terrestrial orchid Bletilla striata

The levels of fruit set in different habitats (three inland populations vs. three coastal populations) and the breeding system in the terrestrial orchid Bletilla striata were examined in two groups of populations in Haenam-gun, Jollanam-do Province in southern Korea. In total inland populations (mean fruit set=13.3%) showed about eight-fold higher fruit set per plant than the coastal populations (1.6%), probably due to mild climatic conditions in the inland populations during the flowering period. Honey bee workers Apis mellifera were observed in the inland populations, but no visitors were found in the coastal populations. However, a great increase in fruit set was achieved by hand-pollination: the percentage of fruit set through artificial self-pollination and geitonogamous pollination were close to 90%, indicating that B. striata is self-compatible. Agamospermy and spontaneous autogamy (automatic self-pollination) were not detected in this study. These indicate that pollinia vectors are essential for achieving fruit set in natural populations of B. striata. Except in one inland population examined in 2001, there was no significant correlation between the patch size per population and the average percentage of fruit set, which might reflect deception-pollination of B. striata and/or paucity of pollinators. Overall levels of fruit sets within and among patches in each population were homogeneous during a two-year period.     

Floral resources,pollinators and fruiting in a threatened tropical deciduous tree

Aims Nothapodytes nimmoniana (Family Icacinaceae) is a deciduous tree species distributed in Asia facing severe population decline. Wood chips from the tree are a source of camptothecin, a globally sought-after alkaloid with cancer-treating properties, and are harvested unsustainably in natural forests. We studied the pollination ecology of the species and asked if there are constraints in pollination and fruiting success in its natural populations. We also discuss the potential effects of wood extraction on pollinators and reproductive success in the population.Methods The study was carried out during the flowering season in two populations with varying exploitation levels, both located in the Protected Area Network in the Western Ghats of Karnataka State. We assessed floral resource availability and measured pollinator diversity and activity in the canopy from the perspective of the biology of the species. We quantified pollinator abundance and percent fruit set as a function of population density.Important findings Flowers belong to the fly pollination syndrome and are open to visits by generalists such as Apis dorsata and Trigona iridipennis. Fruiting success did not vary between exploited and unexploited sites, but there was a significant difference in pollinator assemblage. In a lean male flowering year, fruit set was low, suggesting pollen limitation in the population. No fruiting was recorded in the second year at the site where trees were felled soon after the flowering period. We argue that when male floral resources are altered in exploited populations, pollinators of generalist nature may show a shift in foraging pattern. Also, generalist plants may be as susceptible to pollination loss as are specialist plants. The need for outcrossing in the species would probably exacerbate this vulnerability.     

Floral morph composition and pollen limitation in the seed set of <Emphasis Type="Italic">Nymphoides indica</Emphasis> populations

We investigated the floral morph composition and seed set of the populations in a heterostylous aquatic plant, Nymphoides indica O. Kuntze (Menyanthaceae). The bias of floral morph ratio in the populations was negatively correlated with the seed set. In the populations with biased floral morph ratio, the pollen on the stigma of the common morphs were dominated by pollen of the same morph, resulting in reduced seed set. The addition of pollen of the opposite morph on the stigmas of the common morph resulted in an increase of fruit and seed set rates. The intermorph pollination rate and the fruit and seed set rates of Nymphoides indica were highest when two floral morphs were close in the populations. Various insect species including Lepidoptera, Diptera and Hymenoptera visited the flowers. However, geitonogamous pollination was common because of the short flying distance of the pollinators. These results indicate that the seed set of Nymphoides indica is limited by the shortage of compatible pollens and that the success of intermorph pollination is influenced by spatial distribution of the two floral morphs. As Nymphoides indica is endangered, conservation of the populations with high fecundity is proposed by us.