Stylar polymorphism on the edge: unusual flower traits in Moroccan Narcissus broussonetii (Amaryllidaceae)

Heterostyly and related polymorphisms (e.g. stigma‐height dimorphism) have been used as model systems for studying the origin and maintenance of plant population variability. Stigma‐height dimorphism frequently occurs in Narcissus and is associated with a particular flower shape. In the present study, we describe a new, peculiar case of stigma‐height dimorphism in Narcissus broussonetii, a species on the margin of the geographical distribution of the genus. We determined the stylar condition of N. broussonetii and its variation across populations, analyzed perianth morphology and its relationship with stylar variation, and compared this species with other stylar dimorphic species of the genus. We also studied the incompatibility system and pollination ecology of the species. Narcissus broussonetii is a style‐dimorphic species, as suggested in early studies that were subsequently neglected, and displays unusual flower morphology, with a long floral tube and a virtual absence of a corona. The species shows a late‐ (ovarian) acting incompatibility system and crosses within and between morphs are fertile. We observed short‐tongued diurnal and long‐tongued nocturnal pollinators. Our findings confirm that the presence of the observed dimorphism across populations is most probably the result of the joint action of a nonheteromorphic incompatibility system, extremely long and narrow floral tubes, and a combined role of short‐ and long‐tongued pollinators. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 644–656.     

THE EVOLUTION OF DISTYLY: POLLEN TRANSFER IN ARTIFICIAL FLOWERS

A recent model by Lloyd and Webb derives conditions necessary for the evolution of distyly based on pollen-transfer probabilities between ancestral morphs and invading mutants. We used bumblebees visiting artificial flowers to measure the parameters of the model. Our findings supported the first evolutionary step proposed by the model, establishment of a stigma-height polymorphism. Conditions for the subsequent establishment of an anther-height polymorphism were not satisfied by pollen-transfer patterns alone. Because conditions for the first step are considered more onerous, however, and because the second stage depends on inbreeding depression as well as pollen-transfer patterns, we interpret our results as supporting the plausibility of the Lloyd-Webb model. Video images of bees visiting glass-sided artificial flowers demonstrate a mechanism for disassortative pollination between the ancestral and mutant morph. In general, pollen-transfer probabilities were negatively correlated with the height difference between anthers of the donor and stigma of the recipient. Style length affects bee feeding posture in such a way that disassortative pollination could feasibly occur in the absence of an anther-height polymorphism.     

New perspectives on the evolution of plant mating systems

BACKGROUND: The remarkable diversity of mating patterns and sexual systems in flowering plants has fascinated evolutionary biologists for more than a century. Enduring questions about this topic include why sexual polymorphisms have evolved independently in over 100 plant families, and why proportions of self- and cross-fertilization often vary dramatically within and among populations. Important new insights concerning the evolutionary dynamics of plant mating systems have built upon a strong foundation of theoretical models and innovative field and laboratory experiments. However, as the pace of advancement in this field has accelerated, it has become increasingly difficult for researchers to follow developments outside their primary area of research expertise. SCOPE: In this Viewpoint paper we highlight three important themes that span and integrate different subdisciplines: the changes in morphology, phenology, and physiology that accompany the transition to selfing; the evolutionary consequences of pollen pool diversity in flowering plants; and the evolutionary dynamics of sexual polymorphisms. We also highlight recent developments in molecular techniques that will facilitate more efficient and cost-effective study of mating patterns in large natural populations, research on the dynamics of pollen transport, and investigations on the genetic basis of sexual polymorphisms. This Viewpoint also serves as the introduction to a Special Issue on the Evolution of Plant Mating Systems. The 15 papers in this special issue provide inspiring examples of recent discoveries, and glimpses of exciting developments yet to come.     

Variation and evolution of herkogamy in Exochaenium (Gentianaceae): implications for the evolution of distyly

Backgrounds and Aims

The spatial separation of stigmas and anthers (herkogamy) in flowering plants functions to reduce self-pollination and avoid interference between pollen dispersal and receipt. Little is known about the evolutionary relationships among the three main forms of herkogamy – approach, reverse and reciprocal herkogamy (distyly) – or about transitions to and from a non-herkogamous condition. This problem was examined in Exochaenium (Gentianaceae), a genus of African herbs that exhibits considerable variation in floral morphology, including the three forms of herkogamy.

Methods

Using maximum parsimony and maximum likelihood methods, the evolutionary history of herkogamic and non-herkogamic conditions was reconstructed from a molecular phylogeny of 15 species of Exochaenium and four outgroup taxa, based on three chloroplast regions, the nuclear ribosomal internal transcribed spacer (ITS1 and 2) and the 5·8S gene. Ancestral character states were determined and the reconstructions were used to evaluate competing models for the origin of reciprocal herkogamy.

Key results

Reciprocal herkogamy originated once in Exochaenium from an ancestor with approach herkogamy. Reverse herkogamy and the non-herkogamic condition homostyly were derived from heterostyly. Distylous species possessed pendent, slightly zygomorphic flowers, and the single transition to reverse herkogamy was associated with the hawkmoth pollination syndrome. Reductions in flower size characterized three of four independent transitions from reciprocal herkogamy to homostyly.

Conclusions

The results support Lloyd and Webb''s model in which distyly originated from an ancestor with approach herkogamy. They also demonstrate the lability of sex organ deployment and implicate pollinators, or their absence, as playing an important role in driving transitions among herkogamic and non-herkogamic conditions.     

A one‐locus model of androdioecy with two homomorphic self‐incompatibility groups: expected vs. observed male frequencies

Androdioecy, the occurrence of males and hermaphrodites in a single population, is a rare breeding system because the conditions for maintenance of males are restrictive. In the androdioecious shrub Phillyrea angustifolia, high male frequencies are observed in some populations. The species has a sporophytic self‐incompatibility (SI) system with two self‐incompatibility groups, which ensures that two groups of hermaphrodites can each mate only with the other group, whereas males can fertilize hermaphrodites of both groups. Here, we analyse a population genetic model to investigate the dynamics of such an androdioecious species, assuming that self‐incompatibility and sex phenotypes are determined by a single locus. Our model confirms a previous prediction that a slight reproductive advantage of males relative to hermaphrodites allows the maintenance of males at high equilibrium frequencies. The model predicts different equilibria between hermaphrodites of the two SI groups and males, depending on the male advantage, the initial composition of the population and the population size, whose effect is studied through stochastic simulations. Although the model can generate high male frequencies, observed frequencies are considerably higher than the model predicts. We finally discuss how this model may help explain the large male frequency variation observed in other androdioecious species of Oleaceae: some species show only androdioecious populations, as P. angustifolia, whereas others show populations either completely hermaphrodite or androdioecious.     

Morph-specific variation of floral traits associated with reciprocal herkogamy in natural populations of Primula vulgaris and Primula veris

Documenting the morph-specific variation of floral traits associated with reciprocal herkogamy is of special importance for revealing the functional significance of traits in the evolution and maintenance of the heterostylous syndrome. In order to describe the extent and specificity of variation, stigma height, anther height, stigma-anther separation and corolla tube length were measured on 800 flowers collected in two natural populations of Primula vulgaris and P. veris. Beside the almost complete separation of stigma heights between the two morphs, we found appreciable intermorph overlap in anther height and relatively broad range of stigma-anther separation so far reported for heterostylous species. The corolla tube length–stigma-anther separation relationship showed striking difference between the two floral morphs, which supports the hypothesis that length of the corolla tube plays a more important role in positioning the sex-organs in the appropriate distance in the short-styled than in the long-styled morph.     

Effects of population size on reproductive success of the endangered and endemic species Primula merrilliana

The reproductive ability related to the population size of the endangered and endemic species Primula merrilliana Schltr.was investigated.In 26 populations observed,only four contain more than 500 flowering individuals,whereas most of them(53.8%) consist of less than 100 flowering individuals.Though the ratio of pin and thrum plants keeps its balance at 1:1 for all populations,the frequency of pin and thrum flowers was significantly biased in most small populations.Population size strongly affected reproductive success; plants in small populations produced significantly fewer flowers and fruits per plant and fewer seeds per fruit,and therefore fewer seeds per plant.The floral density was another important factor that influenced the reproductive success of P.merrilliana,because four main reproductive success parameters (fruits per plant,seeds per fruit,seeds per plant,and the proportion of flowers setting fruit) were all positively correlated with floral density.The size of plants and the number of leaves per plant (measure of habitat quality) were not influenced by the variation of population size,suggesting that the reduced fecundity in small populations may not be a consequence of lower habitat quality,inbreeding depression and pollen limitation as a result of less attractiveness in small populations are therefore likely explanations for these patterns.