异型花柱植物喀什补血草的传粉者功能群与花粉转移效率

异型花柱是受遗传控制的花柱多态现象, 被达尔文认为是植物通过在传粉者体表不同部位滞落花粉以促进型间花粉准确传递的一种适应。该现象虽已受到广泛关注, 但在一些花型变异较大且不稳定的传粉系统中, 不同传粉者对各花型繁殖所产生的影响仍知之甚少。该研究以分布于新疆天山南坡的一个有同长花柱共存的异型花柱植物喀什补血草(Limonium kaschgaricum)种群为研究对象, 对其花型构成及频率、传粉者及花粉转移效率等进行了调查分析。结果表明: 1)种群中除了存在雌/雄蕊长度交互对应的长(L)/短(S)花柱型花外, 还有雌/雄蕊同长的花(H型), 且各花型花的花冠口直径、花冠筒长及花粉量等参数间无差异, 但花粉纹饰和柱头乳突细胞形态具二型性。其中, H型花的花粉和柱头形态与L型花(或S型花)的一致。2)花型内和自花授粉均不亲和; 型间授粉时, 花粉和柱头形态不同的花型间亲和, 反之不亲和。3)种群内存在长/短吻两类传粉昆虫。在以短吻传粉者为主的盛花初、中期, L和H型花柱头上的异型花粉数均显著高于S型花的, 且L和S型花高位性器官间的异型花粉传递效率高于低位性器官间的; 而在以长吻传粉者为主的盛花后期, L和S型花的柱头间异型花粉数无显著差异, 且高/低位性器官间具有相同的异型花粉转移效率; 与传粉者出现时期相对应的、在花期不同阶段开放花的结实率也明显不同。4)长/短吻昆虫具明显不同的传粉功能, 短吻昆虫只能对L和H型花进行有效传粉, 且访花频率和型间花粉转移效率较低, 为低效传粉者; 而长吻昆虫对各花型均能有效传粉, 具高的访花频率和型间花粉转移效率, 为高效传粉者。因为长吻昆虫的阶段性出现所形成的不稳定传粉系统, 使低效的短吻昆虫可能会成为种群中花型变异的驱动力, 并使S型花受到更大的选择压力。H型花克服了柱头缩入的弊端, 可能会成为不稳定传粉系统下的一个替代花型而持续存在。     

Pollinator shifts and the loss of style polymorphism in Narcissus papyraceus (Amaryllidaceae)

Darwin proposed that the driving force for the evolution of style polymorphisms is the promotion of cross‐pollination between style morphs, through accurate placement of pollen on the pollinator’s body. This hypothesis has received much attention, but the effect of different pollinators in the fitness of morphs remains poorly understood. Narcissus papyraceus is a style dimorphic species (long ‐L‐ and short ‐S‐ styled) with isoplethic (1 : 1) and L‐monomorphic populations, mainly visited by long‐tongued (LT) nocturnal and short‐tongued (ST) diurnal pollinators, respectively. We studied natural female fertility of morphs, and assessed the role of diurnal and nocturnal pollinators. We also quantified female fertility of the morphs in experimental populations with different morph ratio, exposed to predominately long‐ or short‐tongued pollinators. We found that with LT pollinators, both morphs were successfully pollinated in all morph ratio conditions, suggesting that these insects could be involved in maintenance of the polymorphism, although other factors may also play a role. However, with ST pollinators, S‐plants displayed less fertility than L‐plants, and mating among L‐plants was favoured, implying that the polymorphism is lost. These results underscore the role of pollinators on variations in style polymorphism.     

Pollination ecology in the narrow endemic winter-flowering Primula allionii (Primulaceae)

Reduction of pollen flow can affect plant abundance and population viability and cause selection on plant mating system and floral traits. Little is known on the effect of this phenomenon in species naturally restricted to small and isolated habitats, that may have developed strategies to cope with long-term isolation and small population size. We investigated the pollination ecology of the endemic distylous winter-flowering P. allionii to verify the possible limitation of female fitness due to reduced pollinator visits. We recorded a higher production of pollen grains in long-styled morph, and a higher seed set in short-styled morph. The high intra-morph variability of sexual organ position may explain the hybridization phenomena allowing and easier intra-morph pollination. The fruit set is constant, although its winter-flowering period might decrease pollen transfer. Nevertheless, the lower competition for pollinators with neighbouring plants and the long-lasting anthesis may offset its reproductive success. Even if our results show no evidence of imminent threats, changes in plant–pollinator interactions might increase inbreeding, resulting in an increased extinction risk.     

High herkogamy but low reciprocity characterizes isoplethic populations of Jasminum malabaricum,a species with stigma‐height dimorphism

• Studies of floral polymorphisms have focused on heterostyly, while stigma‐height dimorphism has received considerably less attention. Few studies have examined the reproductive biology of species with stigma‐height dimorphism to understand how factors influencing mate availability and pollen transfer are related to morph ratios in populations.
• Floral morphological traits, especially herkogamy and reciprocity, pollinator visitation, breeding system and spatiotemporal mate availability, are known to affect inter‐morph pollination and morph ratios in species with stigma‐height dimorphism. In this study, we investigated the presence of stigma‐height dimorphism and estimated morph ratios in four naturally occurring populations of Jasminum malabaricum. We quantified morph‐ and population‐specific differences in the abovementioned factors in these populations to understand the observed morph ratios.
• The positions of anthers and stigmas were characteristic of stigma‐height dimorphism, the first report of this polymorphism in the genus. All study populations were isoplethic, implying equal fitness of both morphs. Herkogamy was higher in the short‐styled morph, while reciprocity was higher between the long‐styled stigma and short‐styled anthers. Long‐ and short‐tongued pollinators were common floral visitors, and we observed no differences between morphs in spatiotemporal mate availability or pollinator visitation. Neither morph exhibited self‐ or heteromorphic incompatibility.
• The short‐styled stigma had lower reciprocity but likely receives sufficient inter‐morph pollen from long‐tongued pollinators, and also by avoiding self‐pollination due to higher herkogamy. These results highlight the importance of sufficient effective pollinators and floral morphological features, particularly herkogamy, in maintaining isoplethy in species with stigma‐height dimorphism.

     

茜草科二型花柱植物滇丁香传粉精确性研究

异型花柱是一种受遗传因素控制的花型多态性现象,其适应意义在于提高不同花型间传粉精确性,从而促进异交。为检验二型花柱植物滇丁香（Luculia pinceana）是如何促进异交并确定有效传粉者的类型,实验统计自然种群中长、短柱型的植株数量,测量花部特征,统计自然种群植株柱头上的型间花粉落置的数量和比例,比较型内、型间花粉的萌发率以及花粉管长度,比较不同人工授粉处理结实情况,观察传粉昆虫类型并测量虫体特征。结果表明（1）滇丁香自然居群中长、短柱型植株的数量没有显著性差异。长柱型的筒深、筒直径、开口直径、雄蕊长和柱头长均显著小于短柱型,而长柱型的叶片长和宽、雌蕊长、花药体积、花粉粒体积、柱头厚均显著大于短柱型。（2）自然种群植株柱头上型间花粉所占比例显著低于型内花粉的比例。（3）型间授粉的花粉萌发率以及花粉管长度均大于型内授粉和混合授粉。（4）型间授粉的结实率显著高于型内授粉的结实率,型内授粉具有一定的结实。（5）夜行性长喙条背天蛾（Cechenena lineosa）是滇丁香的有效传粉者,其吻长和滇丁香的花筒深相适应。本研究表明二型花柱植物滇丁香主要是通过与其花部特征相匹配的长喙天蛾实现有效的传粉,通过促进型间花粉的萌发和花粉管的伸长而有利于型间授粉结实,提高传粉精确性。     

Trends in the reproductive biology of Venezuelan Melochia (Malvaceae) species

The breeding system, floral morphology, morph frequency, biomass allocation to floral whorls, pollination and reproductive efficiency were examined in four distylous and three monomorphic Melochia species. Monomorphic species were self-compatible and distylous species were self-incompatible. Flowers of homomorphic species were longer than those of distylous species. Herkogamy was significantly higher in pin than in thrum morphs of the two distylous species, and monomorphic species exhibited the lowest values of herkogamy. Pollen/ovule ratios were similar between monomorphic and dimorphic species, irrespective of the self-incompatibility level. Biomass allocation to flowers was biased toward non-sexual structures, attraction and support. Androecium biomass and androecium/gynoecium biomass ratio were larger in thrum than in pin flowers of distylous species indicating maleness in the thrum morph and femaleness in the pin morph. There was no clear difference between fruit set of monomorphic and dimorphic species; however, the greatest fruit set was found in the monomorphic species, M. pyramidata, which is a self-compatible species. Fruit set was significantly higher in pin than in thrum morphs in three out four distylous species and fecundity was only significantly higher in pin morphs of M. caracasana and M. parvifolia. Melochia species have generalist pollination systems. According to the taxonomic classes and number of pollinator species, M. caracasana and M. parvifolia have similar generalist pollination systems. Our comparative analyses of the characters between floral morphs of distylous species and the relationship with these characters in monomorphic species allow divergences and similarities to be established and different evolutionary trends to be postulated in the breeding systems of Melochia species. Specifically, M. parvifolia and M. caracasana are apparently biased toward monomorphy and dioecy, respectively.     

THE DISSOLUTION OF A COMPLEX GENETIC POLYMORPHISM: THE EVOLUTION OF SELF-FERTILIZATION IN TRISTYLOUS EICHHORNIA PANICULATA (PONTEDERIACEAE)

Eichhornia paniculata (Pontederiaceae) displays a wide range of outcrossing levels as a result of the dissolution of the tristylous genetic polymorphism and the evolution of semihomostyly. Population surveys, comparison of fitness components of the style morphs, and computer simulations were used to investigate the breakdown of tristyly and the selective mechanisms responsible for the evolution of self-fertilization. Of 110 populations surveyed in northeast Brazil and Jamaica, 53% were trimorphic, 25% were dimorphic, and 22% were monomorphic for style morph. The short (S) morph was underrepresented in trimorphic populations and absent from nontrimorphic populations. The mid (M) morph predominated in dimorphic populations and was the only morph in monomorphic populations. Stamen modifications promoting selfing, associated with semihomostyle evolution, were largely confined to the M morph. They were rare in trimorphic populations, common in dimorphic populations, and often fixed in monomorphic populations. Stochastic simulations and comparisons of fruit set in natural populations indicate that founder events, population bottlenecks, and lowered fertility of the S morph due to an absence of long-tongued pollinators can each account for loss of the S morph from trimorphic populations. A reduced level of disassortative mating can accentuate the rate at which the S morph is lost by both random and deterministic processes. Nontrimorphic populations occur at the geographical margins of the region surveyed and tend to be smaller and less dense than trimorphic populations. These observations and the higher fruit set of the M morph relative to the L morph in dimorphic populations suggest that reproductive assurance, favoring selfing variants of the M morph under conditions of low pollinator service, has been of primary importance in the origin of most monomorphic populations. Where pollinator service is reliable, however, automatic selection of selfing genes, aided by mating asymmetries between the morphs, can cause the M morph to spread to fixation in dimorphic populations.     

VARIATION IN THE MATING SYSTEM OF EICHHORNIA PANICULATA (SPRENG.) SOLMS. (PONTEDERIACEAE)

A multilocus procedure was used to estimate outcrossing rates from allozyme data in nine populations of Eichhornia paniculata from NE Brazil and Jamaica. The populations were chosen to represent stages in a proposed model of the evolutionary breakdown of tristyly to semi-homostyly; they differed in morph structure (trimorphic, dimorphic, or monomorphic) and floral traits likely to influence the mating system. The interpopulation range in outcrossing rate, t, was 0.96–0.29. Two additional populations from Jamaica, composed exclusively of self-pollinating, semi-homostylous, mid-styled plants, were invariant at 21 isozyme loci, precluding estimates of outcrossing frequency. Trimorphic populations from Brazil had uniformly high outcrossing rates of 0.96–0.88. Values for the floral morphs within populations were not significantly different. A controlled pollination experiment, comparing the competitive ability of self and cross pollen using the Got-3 marker locus, provided evidence that the maintenance of high outcrossing rates in trimorphic populations results from the prepotency of cross pollen and/or the selective abortion of selfed zygotes. Morph-dependent variation in t was detected within a dimorphic population with the L morph outcrossing with a frequency of 0.76 in comparison with 0.36 in the M morph. The difference in the mating system of floral morphs results from modifications in position of short-level stamens in flowers of the M morph resulting in automatic self-pollination. The occurrence of E. paniculata populations composed exclusively of self-pollinating, mid-styled variants is thought to be associated with the spread of genes modifying stamen position. The high level of self-fertilization demonstrated in the M morph would allow automatic selection of these genes, augmented by fertility assurance in the absence of specialized pollinators.     

The potential for selection on pollen colour dimorphisms in Nigella degenii: morph-specific differences in pollinator visitation, fertilisation success and siring ability

Two subspecies of Nigella degenii (Ranunculaceae) possess a dimorphism in pollen colour and vary extensively in frequency of the two morphs in natural populations. Here we investigate the role of selection on pollen colour during the pollination phase in the two subspecies and its potential contribution to the maintenance of this colour variation. In a combination of common garden experiments and field observations, we obtained data on pollinator visitation rates and explored the effect of pollen colour on fertilisation success and siring ability under conditions of low vs. high pollen competition. In experimental gardens, naïve pollinators responded differently to plants with different pollen colour, but the favoured morph varied between dates and locations, and colour morphs were not visited in a frequency-dependent manner. Donor plants with dark pollen had a reproductive advantage (higher seed set) in single-donor pollinations, but the realised siring ability (measured by progeny morph ratio) was highly variable between different two-donor crosses with no general bias towards the light or dark morph. Therefore, although the dark pollen type appears to have a general selective advantage in terms of fertilisation success, our data are also consistent with a scenario involving the maintenance of both colour morphs, particularly under conditions of high pollen competition, a variable genetic background and/or spatial or temporal variation in the pollinator fauna.     

Identification of effective pollinators of Primula sieboldii E. Morren in a wild habitat in Hiroshima,Japan

Primula sieboldii E. Morren is a heterostylous clonal herb that is widely distributed in Japan but in danger of extinction in the wild. The existence of pollinators in each habitat is imperative for its long-term survival, because seeds can be produced only by insect cross-pollination between different flower morphs. In this study, we identified the pollinators of P. sieboldii in a wild habitat in Hiroshima as those insects that we observed to (a) put the proboscis into a corolla tube, (b) deposit pollen grains on the proboscis, and (c) have a proboscis of appropriate length and width. Effective pollinators were identified from their contribution to pollination. In 2015 and 2016, the flower visitations of 232 and 558 insects, respectively, were recorded and 85 and 13 insects were captured. Two Bombylius species, B. major L. and B. shibakawae Matsumura, accounted for 90% of flower-visiting insects in both years. All 14 species that we captured were considered pollinators of P. sieboldii, because they had proboscises that were long enough to reach pollen and that had pollen grains deposited on them. The total visitation rate of “Bombyliidae” was the highest among all pollinator categories. The results of potential pollen transport per flower per hour, which was based on total pollen number and total visitation rate of each pollinator category, indicated that “Bombyliidae” species were the most effective pollinators of P. sieboldii in this habitat.     

Stigma�Canther reciprocity, pollinators, and pollen transfer efficiency in populations of heterostylous species of Lithodora and Glandora (Boraginaceae)

According to Darwin, the reciprocal position of sexual whorls in heterostylous plants enhances disassortative pollen transfer between different floral morphs. It is believed that greater reciprocity between morphs will promote more efficient transfer of pollen. Additionally, efficient pollination will act as a selective force in achieving greater reciprocity between floral morphs. In this study we test whether variation in reciprocity of sexual organs between morphs is related to the efficiency of pollinators in transferring pollen between them. To do this, we first describe the pollinator??s array in several populations of species of the genus formerly known as Lithodora, which have different types of stylar polymorphism and degrees of reciprocity, and determine their abundance, plant visitation rate, number of flowers visited per plant and handling time in the population. We estimate the efficiency of the pollinator arrays by use of an approximation based on qualitative (location of pollen loads on different areas of insect bodies) and quantitative (plant visitation rate) measurements. Our results show a correlation between the degree of reciprocity and the efficiency of pollinators associated with the populations. These observations suggest that pollinators are a possible selective force driving the evolution of heterostyly.     

Frequency-dependent variation in reproductive success in Narcissus: implications for the maintenance of stigma-height dimorphism

Negative frequency-dependent selection is a major selective force maintaining sexual polymorphisms. However, empirical demonstrations of frequency-dependent reproductive success are rare, particularly in plants. We investigate this problem by manipulating the frequencies of style morphs in a natural population of Narcissus assoanus, a self-incompatible herb with style-length dimorphism and intra-morph compatibility. We predicted that the reproductive success of morphs would vary negatively with their frequency because of the effects of morph-specific differences in sex-organ position on patterns of pollen transfer. This prediction was generally supported. The fruit and seed set of the two morphs did not differ significantly in plots with 1 : 1 morph ratios. However, short-styled plants produced significantly fewer seeds than long-styled plants in monomorphic plots, and significantly more seeds than long-styled plants in plots with 'long-biased' morph ratios. These patterns indicate that in the absence of physiological barriers to intra-morph mating, negative frequency-dependent selection contributes to the maintenance of stylar polymorphism through inter-morph pollen transfer. Our experimental results also provide insights into the mechanisms governing the biased style-morph ratios in populations of Narcissus species.     

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.     

Pollinator response to flower color polymorphism and floral display in a plant with a single-locus floral color polymorphism: consequences for plant reproduction

Variation in flower color, particularly polymorphism, in which two or more different flower color phenotypes occur in the same population or species, may be affected or maintained by mechanisms that depend on pollinators. Furthermore, variation in floral display may affect pollinator response and plant reproductive success through changes in pollinator visitation and availability of compatible pollen. To asses if flower color polymorphism and floral display influences pollinator preferences and movements within and among plants and fitness-related variables we used the self-incompatible species Cosmos bipinnatus Cav. (Asteraceae), a model system with single-locus flower color polymorphism that comprises three morphs: white (recessive homozygous), pink (heterozygous co-dominate), and purple (dominant homozygous) flowers. We measured the preferences of pollinators for each morph and constancy index for each pollinator species, pollination visitation rate, floral traits, and female fitness measures. Flower color morphs differed in floral trait measures and seed production. Pollinators foraged nonrandomly with respect to flower color. The most frequent morph, the pink morph, was the most visited and pollinators exhibited the highest constancy for this morph. Moreover, this morph exhibited the highest female fitness. Pollinators responded strongly to floral display size, while probed more capitulums from plants with large total display sizes, they left a great proportion of them unvisited. Furthermore, total pollinator visitation showed a positive relation with female fitness. Results suggest that although pollinators preferred the heterozygous morph, they alternate indiscriminately among morphs making this polymorphism stable.     

Pollination biology and the impact of floral display, pollen donors, and distyly on seed production in Arcytophyllum lavarum (Rubiaceae)

In animal-pollinated plants, two factors affecting pollen flow and seed production are changes in floral display and the availability of compatible mates. Changes in floral display may affect the number of pollinator visits and the availability of compatible mates will affect the probability of legitimate pollination and seed production. Distyly is a floral polymorphism where long-styled (pin) and short-styled (thrum) floral morphs occur among different individuals. Distylous plants frequently exhibit self and intra-morph incompatibility. Therefore changes in morph abundance directly affect the arrival of compatible pollen to the stigmas. Floral morph by itself may also affect female reproductive success because floral morphs may display differences in seed production. We explored the effects of floral display, availability of neighboring compatible mates, and floral morph on seed production in the distylous herb ARCYTOPHYLLUM LAVARUM. We found that floral display does not affect the mean number of seeds produced per flower. There is also no effect of the proportion of neighboring legitimate pollen donors on seed production in pin or thrum flowers. However, floral morphs differed in their female reproductive success and the thrum morph produced more seeds. Hand pollination experiments suggest that differences in seed production between morphs are the result of pollen limitation. Future research will elucidate if the higher seed production in thrum flowers is a consequence of higher availability of pollen donors in the population, or higher efficiency of the pin morph as pollen donor.     

STIGMATIC POLLEN LOADS IN POPULATIONS OF PONTEDERIA CORDATA FROM THE SOUTHERN U.S.

Stigmatic pollen loads were examined in four tristylous populations of Pontederia cordata from the southern U.S. to evaluate Darwin's hypothesis that floral trimorphism promotes legitimate pollination. In each population, morph frequency, pollen production, the composition of stigmatic pollen loads and seed production were estimated. Goodness-of-fit tests and a measure of the efficiency of legitimate pollination were employed to evaluate pollen load patterns. Results from the four populations demonstrate that the long-styled (L) morph usually experiences legitimate pollination, whereas the mid-styled (M) and short-styled (S) morphs often display random pollination. However, at Taylor Road (Louisiana) all three floral morphs exhibited statistically significant levels of legitimate pollination. The size of legitimate pollen loads suggests that in populations of P. cordata tristyly may have only a minor influence on the fecundity of morphs. Two hypotheses are proposed to explain the maintenance of floral trimorphism in contemporary populations of Pontederia spp. The first assumes the polymorphism is selectively neutral; the second suggests that it contributes towards the male component of fitness by increasing the distance, and hence the number of genets, that pollen can be transported to by pollinators. In clonal species, such as Pontederia cordata, where geitonogamous pollinations are promoted by large floral displays, floral mechanisms that increase pollen carryover may have selective value.     

Female reproductive success in gynodioecious Thymus vulgaris: pollen versus nutrient limitation and pollinator foraging behaviour

Gynodioecy is a dimorphic breeding system in which female individuals coexist with hermaphroditic individuals in the same population. Females only contribute to the next generation via ovules, and many studies have shown that they are usually less attractive than hermaphrodites to pollinators. Several mechanisms have been proposed to explain how females manage to persist in populations despite these disadvantages. The ‘resource reallocation hypothesis’ (RRH) states that females channel resources not invested in pollen production and floral advertisement towards the production of more and/or larger seeds. We investigated pollination patterns and tested the RRH in a population of Thymus vulgaris. We measured flower display, flower size, nectar production, visitation rates, pollinator constancy and flower lifespan in the two morphs. In addition, we measured experimentally the effects of pollen and resource addition on female reproductive success (fruit set, seed set, seed weight) of the two morphs. Despite lower investment in floral advertisement, female individuals were no less attractive to pollinators than hermaphrodites on a per flower basis. Other measures of pollinator behaviour (number of flowers visited per plant, morph preference and morph constancy) also showed that pollinators did not discriminate against female flowers. In addition, stigma receptivity was longer in female flowers. Accordingly, and contrary to most studies on gynodioecious species, reproductive success of females was not pollen limited. Instead, seed production was pollen limited in hermaphrodites, suggesting low levels of cross‐pollination in hermaphrodites. Seed production was resource limited in hermaphrodites, but not in females, thus providing support for the RRH. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 395–408.     

Asymmetrical mating patterns and the evolution of biased style-morph ratios in a tristylous daffodil.

Non-random mating in plant populations can be influenced by numerous reproductive and demographic factors, including floral morphology and inter-plant distance. Here, we investigate patterns of outcrossed mating through male function in Narcissus triandrus, a tristylous, bee-pollinated wild daffodil from the Iberian Peninsula, to test pollen transfer models which predict that floral morphology promotes asymmetrical mating and biased morph ratios. Unlike other tristylous species, N. triandrus has an incompatibility system that permits intra-morph mating and long-level rather than mid-level stamens in the L-morph. Incomplete sex-organ reciprocity should result in significant intra-morph mating in the L-morph. We measured mating patterns in two L-biased populations--dimorphic (two style morphs) and trimorphic (three style morphs)--using multilocus genotyping and maximum-likelihood-based paternity analysis. We also examined the spatial distribution of style morphs and neutral markers to investigate the potential consequence of spatially restricted mating on morph ratios. As predicted, we detected significant amounts of intra-morph mating in the L-morph in both populations. Pollen transfer coefficients generally supported predictions based on the Darwinian hypothesis that anthers and stigmas of equivalent level promote pollinator-mediated cross-pollination in heterostylous populations. There was evidence of significant spatial aggregation of both style morphs and neutral markers in populations of N. triandrus, probably as a result of restricted pollen and seed dispersal. Our results provide empirical support for theoretical models of pollen transfer, which indicate that the commonly observed L-biased morph ratios in Narcissus species result from significant intra-morph mating in the L-morph because of its atypical floral morphology.     

Pollinator effectiveness and fruit set in common ivy, Hedera helix (Araliaceae)

Flowers of common ivy (Hedera helix L.) provide late season pollen and nectar for several insect groups, and its fruits are a winter and spring food source for frugivorous birds. Ivy benefits from insect pollination in order to set fruit, but it is unknown which flower-visiting insects are the most effective pollinators. Our observations suggest that Vespula wasps are potentially the most effective pollinators since they were frequent visitors, had relatively fast foraging rates, carried large numbers of pollen grains on their bodies and had the highest ‘Pollination potential PP index’ score (a measure of pollinator effectiveness) of all the insect groups examined. There was also a positive linear relationship between the proportion of ivy flowers that set fruit and wasp foraging activity in 0.5 m² quadrats. Visits by Vespula wasps may therefore be important for ensuring a supply of ivy fruits for birds.     

Population size, pollination and reproductive success in two endangered Genista species

Habitat fragmentation often leads to small and isolated plant populations as well as decreased habitat quality. These processes can fundamentally disrupt the interactions between plants and pollinators and decrease reproductive success. This concerns especially self-incompatible, non-clonal species that depend on pollination for successful reproduction.In two rare and endangered heathland plant species, Genista anglica and G. pilosa, we examined pollination and reproduction in relation to population size. Eight populations of G. anglica and ten populations of G. pilosa were surveyed in the vicinity of Bremen, NW-Germany. We counted the visits of pollinators (honeybees, bumblebees, and other insects) and determined the reproductive output of the observed shoots.Contrary to our expectation to find increased pollinator visitation rates in larger populations of both Genista species, the number of flower-visiting insects was unrelated to the number of flowering shoots. Increasing shoot length had a positive and increasing temperature a negative impact on the number of visiting honeybees and bumblebees. Despite the general absence of population size effects on pollinator numbers, the number of fruits and seeds in G. anglica increased with increasing population size. Fruit and seed set in G. pilosa were negatively related to the number of ‘other insects’. Our field observations showed that larger populations of both Genista species flowered earlier than smaller populations and much earlier than reported in the literature. Flowering in large populations therefore tends to coincide less well with pollinator abundance, and this may cause a disruption of the temporal coincidence between flowering phenology and pollinator activity.