Style Morphs Research Articles

The enantiostylous floral polymorphism of Barberetta aurea (Haemodoraceae) facilitates wing pollination by syrphid flies.

Sexual polymorphisms of flowers have traditionally been interpreted as devices that promote cross-pollination, but they may also represent adaptations for exploiting particular pollination niches in local environments. The cross-pollination function of enantiostyly, characterized by flowers having either left- or right-deflected styles, has been uncertain in some lineages, such as the Haemodoraceae, because the positioning of stamens and styles is not always completely reciprocal among morphs. We examined the floral biology of populations of the poorly known species Barberetta aurea (Haemodoraceae) across its native range in South Africa to establish the general features of its enanatiostylous reproductive system and the agents and mechanism of pollen transfer. We confirmed that B. aurea has a system of dimorphic enantiostyly. Style morph ratios varied among populations sampled, but with an overall tendency to being equal. Crossing experiments demonstrated that B. aurea is fully self-compatible, that intra- and inter-morph crosses are equally fertile and that it is wholly dependent on pollinator visits for seed production. Pollination is mainly by syrphid flies that transfer the sticky pollen via their wings, which contact the anthers and stigma precisely as they hover during approach and feeding. The majority of syrphid fly visitors feed on a film of highly concentrated nectar situated at the base of ultraviolet-absorbent 'nectar guides'. Because one of the three stamens is deflected in the same direction as the style, we predicted a high likelihood of intra-morph pollination, and this was corroborated by patterns of transfer of coloured dye particles in cage experiments involving syrphid flies. Barbaretta aurea exhibits dimorphic enantiostyly and, in contrast to most enantiostylous species, which are pollinated by bees, its flowers are specialized for pollination by syrphid flies. The lack of complete reciprocity of the enantiostylous arrangement of sexual organs facilitates both inter- and intra-morph pollen transfer on the wings of these flies.

Heterostyly, incompatibility system and pollinators in Varronia spinescens Borhidi (L.) (Cordiaceae)

Heterostyly, the reciprocal position of sexual organs between different morphs within a population, is assumed to promote cross-pollination in hermaphroditic plants. Heterostyly is usually linked to an incompatibility system where only the crosses between the different morphs produce fruit. Although heterostyly has been frequently studied in plant species with specialized pollination systems, few studies have investigated this in species with a broad spectrum of floral visitors. Here, we describe the heterostyly of Varronia spinescens and conducted hand-pollination experiments in the field to determine the incompatibility system. We also analysed perianth morphology, morph-ratio, adaptive inaccuracy in reciprocity and pollinator visitation rate in relation to floral morph and number of open flowers. Our results indicate that V. spinescens is a distylous species with a high degree of reciprocity between the sexual organs, presenting a non isoplethic morph ratio with a bias towards the long style (L-) morph. The incompatibility system is heteromorphic and the studied population received visits by a very high number of visitor and pollinator species. Varronia spinescens is thus the heterostylous species of the genus with the second-highest number of registered pollinators. Our results suggest that in a generalist pollination system with little floral morphological restriction to pollinators, the maintenance of sexual polymorphism is likely associated with a high reciprocity between morphs.

Floral Scent Variation in the Heterostylous Species Gelsemium sempervirens.

Gelsemium sempervirens (L.) W.T. Aiton, a distylous woody vine of the family Gelsemiaceae, produces sweetly fragrant flowers that are known for the toxic alkaloids they contain. The composition of this plant’s floral scent has not previously been determined. In this study, the scent profiles of 74 flowers obtained from six different wild and cultivated populations of G. sempervirens were measured by solid phase microextraction-gas chromatography-mass spectrometry (SPME-GC-MS). There were 81 volatile organic compounds identified and characterized as benzenoids, terpenoids, fatty acid derivatives, and yeast associated compounds. The most abundant compound was benzaldehyde (23–80%) followed by ethanol (0.9–17%), benzyl benzoate (2–15%), 4-anisaldehyde (2–11%), (Z)-α-ocimene (0–34%), and α-farnesene (0.1–16%). The impacts of geographic location, population type (wild or cultivated), and style morph (L = long, S = short) on scent profile were investigated. The results showed no relationship between geographic location or population type and volatile organic compounds (VOC) profile, but did show a significant scent profile difference between L and S morphs based on non-metric multidimensional scaling (NMDS) using Bray-Curtis similarity indices. The L morphs contained higher amounts of benzenoids and the S morphs contained higher amounts of terpenoids in their scent profiles. The L morphs also produced a higher total abundance of scent compounds than the S morphs. This study represents the first floral scent determination of G. sempervirens finding significant variation in scent abundance and composition between style morphs.

The genetic architecture of tristyly and its breakdown to self-fertilization.

The floral polymorphism tristyly involves three style morphs with a reciprocal arrangement of stigma and anther heights governed by two diallelic loci (S and M). Tristyly functions to promote cross-pollination, but modifications to stamen position commonly cause transitions to selfing. Here, we integrate whole-genome sequencing and genetic mapping to investigate the genetic architecture of the M locus and the genetic basis of independent transitions to selfing in tristylous Eichhornia paniculata. We crossed independently derived semi-homostylous selfing variants of the long- and mid-styled morph fixed for alternate alleles at the M locus (ssmm and ssMM, respectively), and backcrossed the F1 to the parental ssmm genotype. We phenotyped and genotyped 462 backcross progeny using 1450 genotyping-by-sequencing (GBS) markers and performed composite interval mapping to identify quantitative trait loci (QTL) governing style-length and anther-height variation. A QTL associated with the primary style-morph differences (style length and anther height) mapped to linkage group 5 and spanned ~13-27.5Mbp of assembled sequence. Bulk segregant analysis identified 334 genes containing SNPs potentially linked to the M locus. The stamen modifications characterizing each selfing variant were governed by loci on different linkage groups. Our results provide an important step towards identifying the M locus and demonstrate that transitions to selfing have originated by independent sets of mating-system modifier genes unlinked to the M locus, a pattern inconsistent with a recombinational origin of selfing variants at a putative supergene.

Reproduction: The genetic basis of heterostyly.

Two recent studies revealed the genetic architecture of the long fascinating heterostyly in Primula and that the absence of a CYPT gene determines the long style morph.

Stochastic Processes during Invasion: The Influence of Population Size on Style-Morph Frequency Variation in Lythrum salicaria (Purple Loosestrife)

Premise of research. During biological invasion, the genetic diversity of populations may be reduced by founder events and genetic drift. The floral polymorphism tristyly provides an exceptional opportunity to investigate the influence of stochastic forces on the maintenance of genetic polymorphism, because small population size leads to a characteristic signature of morph loss from populations. Here, we investigate the relations between population size and morph-frequency variation in invasive populations of tristylous Lythrum salicaria in Ontario, Canada. We also compare our results to a similar survey conducted 25 years ago in the same region.Methodology. We surveyed the size and morph ratios of 114 L. salicaria populations in 2013. We calculated the relations between population size and style morph absence, population size and style morph evenness, and the number of populations lacking particular style morphs. For comparison of the patterns of morph-frequency variation between surveys, we used a sampl...

Variation in style morph frequencies in tristylous Lythrum salicaria in the Iberian Peninsula: the role of geographical and demographic factors.

The balance between stochastic forces and negative frequency-dependent selection largely determines style morph frequencies in heterostylous populations. Investigation of morph frequencies at geographical range limits can provide insights into the forces maintaining the floral polymorphism, and the factors causing biased morph ratios. Here, we investigate style morph frequencies in populations at the south-western European range limit of tristylous Lythrum salicaria, to explore the role of demographic and geographical factors influencing morph ratios in its native range. We measured morph composition and evenness, and the size of 96 populations, along a north to south latitudinal transect from Galicia to Andalucia, Iberian Peninsula, traversing a steep climatic gradient. To examine the potential influence of morph-specific fitness components on morph ratios, we examined reproductive traits in 19 populations. Most populations of L. salicaria were trimorphic (94·79 %), the majority exhibiting 1 : 1 : 1 morph ratios (68·75 %). Populations with biased morph ratios had a deficiency of the short-styled morph. Population size and morph evenness were positively associated with latitude, with smaller populations and those with less even morph ratios occurring towards the south. Greater variance in morph evenness was evident at the southern range margin. There were no consistent differences in components of reproductive fitness among style morphs, but southern populations produced less fruit and seed than more northerly populations. Our results demonstrate the influence of finite population size on morph frequencies in L. salicaria. However, they also illustrate the resilience of Iberian populations to the factors causing deviations from isoplethy and morph loss, especially at the southern range limit where populations are smaller. The maintenance of tristyly in small populations of L. salicaria may be aided by the genetic connectivity of populations in agricultural landscapes resulting from gene flow through pollen and seed dispersal.

Postpollination discrimination between self and outcross pollen covaries with the mating system of a self-compatible flowering plant.

Variation in the mating system of hermaphroditic plant populations is determined by interactions between genetic and environmental factors operating via both pre- and postmating processes. Models predicting the maintenance of intermediate outcrossing rates in animal-pollinated plants often assume that the mating system is primarily controlled by floral morphology and pollinator availability, but rarely has the influence of postpollination processes on variation in outcrossing been examined. We investigated the influence of stylar discrimination between illegitimate and legitimate pollen-tube growth and the pollen-load capacity of stigmas on mating-system variation in the annual, tristylous species Eichhornia paniculata using controlled crosses and genetic markers. This species exhibits an exceptionally broad range of outcrossing rates in natural populations. There was significant variation among populations in the pollen-load capacity of stigmas and the ability of styles to discriminate between illegitimate vs. legitimate pollen. There was strong correspondence between stylar-discrimination ability and variation in outcrossing rate among populations and style morphs. The combination of stigmatic pollen-load capacity and stylar discrimination explained more than 80% of the variation in outcrossing rates among populations. The finding that stigmatic pollen-load capacity and stylar-discrimination ability contributed significantly to explaining the wide range of outcrossing rates in E. paniculata suggests that postpollination mechanisms play an important role in governing mating patterns in this species. The difference in levels of stylar discrimination between outcrossing and selfing populations may reflect a trade-off between selection for increased outcrossing and greater reproductive assurance.

Floral variation and environmental heterogeneity in a tristylous clonal aquatic of the Pantanal wetlands of Brazil.

The balance between stochastic forces and frequency-dependent mating largely governs style morph frequencies in heterostylous populations. In clonal species, deviations from equal morph ratios often result from founder events and unfavourable conditions for sexual reproduction. The aim of this study was to investigate whether different flooding regimes, because of their influence on sexual vs. clonal reproduction, are associated with regional variation in morph frequencies and floral trait differentiation in populations of the clonal, tristylous, aquatic Eichhornia azurea (Pontederiaceae) in the Pantanal wetlands of Brazil. Style morph frequencies were sampled from 73 populations distributed across four flooding regimes differing in depth and duration. Measurements of flower size, sex-organ dimension, pollen size and pollen production were made in selected populations, and pollinator assemblages and their functional traits were recorded. Most populations of E. azurea were tristylous (78 %), but the majority exhibited uneven morph ratios. The frequency of the mid-styled morph was significantly lower than that of the long- and short-styled morphs. Morph evenness was positively associated with population size but not with flooding regime. There were significant phenotypic differences among flooding regimes for all floral traits, including populations with reduced flower size, sex-organ length and smaller pollen. Pollinator assemblages varied with flood duration. The similar morph structure and evenness of populations, regardless of flooding regime, suggest that sexual reproduction and clonal dispersal are sufficiently common to prevent the signature of founder events from dominating in a region. However, the pervasive occurrence of biased morph ratios in most populations suggests that many are in a non-equilibrium state. The reduced frequency of the mid-styled morph in trimorphic and dimorphic populations may be associated with the weak self-incompatibility of this morph resulting in selfing and inbreeding depression. Clonality in E. azurea and the weak self-incompatibility of the mid-styled morph may make it more vulnerable to geitonogamous selfing.

Variation in Sex Allocation and Floral Morphology in an Expanding Distylous Plant Hybrid Complex

Premise of research. Sex allocation, the relative energy devoted to producing pollen, ovules, and floral displays, can significantly affect reproductive output and population dynamics. In this study, we investigated floral morphology and gamete production in bisexual, distylous plants from a self-incompatible hybrid complex (Piriqueta cistoides ssp. caroliniana Walter [Arbo]; Turneraceae). Sampling focused on two parent types (C, V) and their stable hybrid derivative (H). Since H morphotypes are heterotic for growth and fruit production, we hypothesized that they would produce larger flowers with more gametes. We also anticipated that plants with long styles (long morphs) would produce less pollen than short morphs, since long-morph pollen is larger.Methodology. Over two consecutive summers, flowers were collected from 1465 individual plants in 28 field populations. Floral parameters were measured digitally, and each flower’s pollen number, ovule number, and stigma-anther separation was quantified under a dissecting microscope. Gamete production (n = 332) and stigma-anther separation (n = 119) were also quantified for plants from a greenhouse accession.Pivotal results. Floral display differed among morphotypes, with H plants producing the largest flowers and C plants displaying the least petal separation. Hybrid morphotypes produced significantly more pollen than parental morphotypes, and pollen quantity was significantly greater for long morphs. Ovule production, however, was greatest for V flowers. Stigma-anther separation differed between years and style morphs (greater for short morphs) but not among morphotypes or within a single season.Conclusions. Differences in pollen production between morphs were not consistent with trade-offs in pollen size and number or selection for increased male function in short morphs. Greater stigma-anther separation in short morphs supported the hypothesis of selection to reduce pollen interference. Enhanced floral display and pollen production followed other heterotic traits observed in H morphotypes. The superior ability of H morphotypes to attract pollinators and sire seeds might partially explain this hybrid zone’s continuing expansion.

Long‐tongued insects promote disassortative pollen transfer in style‐dimorphic Narcissus papyraceus (Amaryllidaceae)

Summary In hermaphroditic flowers, reciprocal herkogamy, for example heterostyly, enhances pollen transfer between floral morphs (disassortative pollination) while avoiding self‐interference between sexual organs. By contrast, disassortative pollination might be compromised in style‐dimorphic flowers, which lack perfect reciprocity between the two floral morphs. This suboptimal functioning has been considered to explain why stylar dimorphism is rare in nature. Some style‐polymorphic species receive a wide array of floral visitors, including long‐tongued insects that feed on nectar and short‐tongued insects that feed on pollen. Differences in the morphology and behaviour of these two insect types could be manifested as different pollination patterns in each floral morph. By observing individual pollinators and pollen deposition and delivery in small field‐based experimental arrays with emasculated and intact flowers, we studied pollination patterns mediated by different insect types (long‐ and short‐tongued) in the two floral morphs (long‐ and short‐styled) of the style‐dimorphic Narcissus papyraceus. We investigated the patterns of pollen transfer between and within style morphs in cross‐pollinations, as well as self‐pollination and pollen removal rates, for each floral morph mediated by each insect type. Long‐tongued insects were efficient pollinators of both floral morphs as they removed little pollen from the anthers but deposited comparatively large amounts on the stigmas. Although disassortative pollen transfer and assortative pollen transfer were equally high to the long‐styled morph, the former prevailed in the short‐styled morph. Short‐tongued insects removed large amounts of pollen from the anthers but deposited only a few pollen grains on the long‐styled stigmas and a negligible number of grains on short‐styled stigmas regardless of the morph of the donor. Synthesis. In this study, we provide empirical support for the hypothesis that, under the action of long‐tongued pollinators, pollen transfer patterns in the stylar‐dimorphic Narcissus papyraceus resemble in part those of heterostylous species. In addition, we found that short‐tongued insects act mostly as pollen thieves, thereby limiting the male fitness of both style morphs, besides depleting the female fitness of S‐morph plants. In view of these results, we propose that the differing pollination efficiencies of floral visitors, in addition to their frequency, are key in determining the morph ratio of populations in this Narcissus.

Genotypic diversity and genetic structure of populations of the distylous aquatic plantNymphoides peltata(Menyanthaceae) in China

AbstractNymphoides peltatais a distylous aquatic plant. In order to explore the relative importance of clonal growth and sexual reproduction within populations and also to reveal the origin of the commonly occurring one style morph populations in China, we studied the clonal diversity and population genetic structure of this species. By using seven single sequence repeat primer pairs, a total of 192 multilocus genotypes (MLGs) were identified among the 472 samples from the 21 populations analyzed; each population had two or more MLGs. The number of MLGs was lower for the one style morph populations than distylous populations. Analysis of molecular variance showed that a smaller proportion of genetic variation resided among populations (36.8%). A neighbor‐joining tree indicated that there was no significant geographical structure of the genetic variation among populations ofN. peltata. The Bottleneck tests indicated that there had been significant bottlenecks in most of the studied populations. The successful sexual recruitment from seeds may have contributed to the high clonal diversity in distylous populations ofN.peltatain China. Demographic stochasticity or bottleneck should account for the one style morph population with several MLGs. And also the bottleneck effects or demographic changes caused by human activities occurring randomly in populations should account for the finding of no significant geographical structure of the genetic variation among the investigated populations ofN.peltata.

DECONSTRUCTING HETEROSTYLY: THE EVOLUTIONARY ROLE OF INCOMPATIBILITY SYSTEM, POLLINATORS, AND FLORAL ARCHITECTURE

Darwin's early work on heterostyly and related style polymorphisms (the presence of two or three style morphs within a population) generated much interest to understand how precise interactions between ecological and genetic mechanisms influence the evolution of floral diversity. Here we tested three key hypotheses proposed to explain the evolution of heterostyly: (i) the presence of self-incompatibility; (ii) the role of pollinators in promoting dissasortative mating; and (iii) floral architecture, which restricts pollinators' movements and ensures more exact pollen deposition on their bodies. We combined data from experiments, field observations, and published studies to test whether evolution of style polymorphism in Narcissus is driven by the incompatibility system, pollinator guilds, or floral architecture, within a phylogenetic framework. Neither differences in pollinator environment nor the presence of genetic self-incompatibility were correlated with presence of style polymorphism. However, our results indicate that the evolution of style polymorphism was driven by the presence of a narrow and long floral tube.

The effect of host abundance on the distribution and impact of biocontrol agents on purple loosestrife (Lythrum salicaria, Lythraceae)

The spread and efficacy of biological control agents are thought to increase with the regional frequency of host populations. We tested this hypothesis by quantifying the occurrence of and damage by 2 biocontrol agents, Galerucella pusilla and G. calmariensis beetles, on 52 introduced stands of purple loosestrife (Lythrum salicaria) in Ontario, Canada. An analysis of herbarium records combined with population-genetic analysis of style morph frequencies and extensive field observations made across Ontario over a 25-y period indicate that L. salicaria populations are more densely distributed in eastern than central Ontario. However, we did not detect the control agents more frequently nor was damage by biocontrol agents higher in 14 eastern stands than 17 central stands. By comparing 20 stands where beetles had been deliberately released with 32 for which there is no recorded history of beetle release, we found that beetles had spread widely beyond documented release sites. However, plants at release sites were more heavily damaged than those at non-release sites, such that previous surveys monitoring only release sites overestimated the efficacy of control. Overall, we detected herbivory by Galerucella spp. in 88% of L. salicaria stands throughout eastern and central Ontario, and only 5% of 5957 plants sampled seem to have entirely escaped damage. We recommend an iterative post-release monitoring study to determine the broader efficacy of the control program using the baseline quantitative data presented here.

The role of short-tongued insects in floral variation across the range of a style-dimorphic plant.

Heterostyly and related style polymorphisms are suitable model systems to evaluate the importance of functional pollinators in the maintenance of population variability. In Narcissus papyraceus different functional pollinators, incompatibility system and flower morphology have been proposed to influence the maintenance of polymorphism through their effect on disassortative mating. Here a test is done to find out if the visitation rate of long- versus short-tongued pollinators correlates with the morph ratio and if the latter is related to other flower traits of the species across its main geographic range. Floral traits from 34 populations in the south-west of the Iberian Peninsula and in north-west Africa were measured, perianth variation was described and a comparison was made of allometric relationships between sex organs and floral tube. Correlations between pollinator guilds, stigma-anther separation of reciprocal morphs (our proxy for disassortative mating) and morph-ratio variation were analysed. Finally, the incompatibility system of the species in the northern and southern borders of its distribution are described. Flowers from southern populations were significantly larger than flowers from centre and northern populations. The abundance of short-styled plants decreased gradually with increasing distance from the core region (the Strait of Gibraltar), with these disappearing only in the northern range. Although there was a significant difference in stigma-anther separation among populations, morph ratio was not associated with reciprocity or floral tube length. Long-style morph frequency increased with short-tongued pollinator visitation rate. Populations from both edges of the distribution range were self-incompatible and within- and between-morph compatible. The style morph ratio changed gradually, whereas perianth trait variation showed abrupt changes with two morphotypes across the range. The positive relationship between the visitation rate of short-tongued pollinators and the decrease of the short-style morph supports our initial hypothesis. The results highlight the importance of different pollinators in determining the presence of style polymorphism.

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.

Variation of Self‐Incompatibility within Invasive Populations of Purple Loosestrife (Lythrum salicariaL.) from Eastern North America

Colonization may favor self‐compatibility (SC) in invasive plants, a process consistent with Baker’s law. We investigated this hypothesis in invasive eastern North American populations of tristylous Lythrum salicaria L. (purple loosestrife) by controlled self‐ and cross‐pollinations of 124 plants sampled from 12 populations grown under uniform glasshouse conditions. We evaluated whether populations at the northern front of the invasion exhibited higher levels of SC than southern populations, which are closer to source populations for the North American invasion. We also sought evidence for morph‐specific differences in the strength of trimorphic incompatibility. We used the ASTER statistical program to jointly model three measures of SC: (1) the probability of producing at least one seed following self‐pollination, (2) the number of fruits produced by self‐pollination, and (3) the average number of seeds per fruit resulting from self‐pollination. Just over one‐quarter of plants set at least one fruit after self‐pollination, and there was a significant effect of style morph on variation in SC, with the mid‐styled morph most compatible. Although variation for SC was detected in 11 of the 12 populations, there was no evidence that SC increased toward the northern range limit as a result of the invasion process.

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.

Population structure and genetic diversity in tristylous Narcissus triandrus: insights from microsatellite and chloroplast DNA variation

We investigated cpDNA sequence and nuclear microsatellite variation among populations of the wild daffodil Narcissus triandrus to examine the role of historical vs. contemporary forces in shaping population structure, morphological differentiation and sexual-system evolution. This wide-ranging heterostylous species of the Iberian Peninsula is largely composed of two allopatric varieties (vars. cernuus and triandrus), and populations with either stylar trimorphism or dimorphism. Dimorphic populations only occur in var. triandrus, are mainly restricted to the northwestern portion of the species range, and uniformly lack the mid-styled morph (M-morph). Chloroplast DNA (cpDNA) sequence variation revealed strong geographical structuring and evidence for a fragmentation event associated with differentiation of the two varieties. In var. triandrus, population fragmentation, restricted gene flow and isolation-by-distance were also inferred. Significant differences in genetic diversity and population structure between the two varieties likely reflect historical and contemporary differences in demography and gene flow among populations. Discordance between cpDNA markers and both microsatellite and morphological variation indicate that hybridization has occurred between the two varieties at contact zones. There were no differences in genetic diversity or population structure between dimorphic and trimorphic populations, and chloroplast haplotypes were not associated with either sexual system, indicating transitions in morph structure within each maternal lineage. M-morph frequencies were positively correlated with differentiation at microsatellite loci, indicating that the evolutionary processes influencing these neutral markers also influence alleles controlling the style morphs.

Distylic Hottonia palustris shows high reproductive success in small populations despite low genetic variability

Hottonia palustris L. (Primulaceae) is characterized by a heteromorphic incompatibility system. The strategy of distylic ramets is believed to promote outcrossing, to maintain overall genetic diversity and to prevent inbreeding depression. In spite of this distyly, an extremely low amount of allozymic diversity was observed in 545 individual ramets from 14 populations in Flanders (Belgium). A possible explanation for such low genetic variation is discussed in relation to the vegetative propagating abilities and the ecological niche width of the species. In contrast to the uniformity in allozymes as well as to the feature of single morph populations, there was a high variability in reproductive success between populations such as the number of seeds per ramet (425–2633), the number of flowers per ramet (9–36) and the mean weight of seeds (0.03–0.17 mg). Small populations and even those consisting of only one style morph may show a high reproductive success. As a whole, H. palustris showed a negative relationship of reproductive success with the surface area of its populations.