The Affinities and Relationships of the Japanese Endemic Pseudopyxis (Rubiaceae-Paederieae)

Abstract Selected character states of the presumed allies of Pseudopyxis are described, compared and discussed. Particular attention is paid to the fruits and diaspores and their ways of development and differentiation. It is presumed that 1) Pseudopyxis and Serissa are closely allied, that 2) these two genera are, in turn, closely related to Leptodermis and the monotypic Spermadictyon (the latter possibly not generically distinct from the former), that 3) Paederia loosely links up to four mentioned genera, and that 4) the monotypic Aitchisonia stands rather isolated on account of various peculiarities.     

Pollen flow in a population of Primula vulgaris Huds.

A British population of Primula vulgaris was visited four times between 1971 and 1976 for study of the nature and functioning of distyly. The stainability and production of pollen from pins exceeded those of thrums. About 20% of pin pollen and 48% of thrum pollen was removed from anthers by insect visitors. Pins and thrums were present in a 1:1 ratio in 1971 but pins outnumbered thrums in 1976. Thrums produced more seeds per flower than pins though the number of ovules produced by each was similar. Two methods for collecting stigmas from open flowers and analyzing them for pollen loans produced somewhat different results. For pin stigmas, the pollen load consisted of 2–23% thrum pollen; for thrum stigmas, the pollen load consisted of 0–71% pin pollen with most stigmas having less than 50% pin pollen. In general, intermorph pollen flow is less than would be expected if pollen flow were random. It is probable that most intramorph pollen on stigmas is a result of self- or geitonogamous pollination. The extensive literature concerning the natural pollinators of the primrose is reviewed. Although Darwin's hypothesis concerning the functional significance of distyly in promoting intermorph pollination was never quantified, the pollen flow patterns observed in P. vulgaris are unexpectedly deviant and are similar to those patterns observed in several unrelated heterostylous species in other plant families.     

Comparison of pollen transfer dynamics by multiple floral visitors: experiments with pollen and fluorescent dye

BACKGROUND AND AIMS: Most plant species are visited by a diversity of floral visitors. Pollen transfer of the four most common pollinating bee species and one nectar-robbing bee of the distylous plant Gelsemium sempervirens were compared. METHODS: Naturally occurring pollen loads carried by the common floral visitor species of G. sempervirens were compared. In addition, dyed pollen donor flowers and sequences of four emasculated recipient flowers in field cages were used to estimate pollen transfer, and the utility of fluorescent dye powder as an analogue for pollen transfer was determined. KEY RESULTS: Xylocopa virginica, Osmia lignaria and Habropoda laboriosa carried the most G. sempervirens pollen on their bodies, followed by Bombus bimaculatus and Apis mellifera. However, B. bimaculatus, O. lignaria and H. laboriosa transferred significantly more pollen than A. mellifera. Nectar-robbing X. virginica transferred the least pollen, even when visiting legitimately. Dye particles were strongly correlated with pollen grains on a stigma, and therefore provide a good analogue for pollen in this system. The ratio of pollen : dye across stigmas was not affected by bee species or interactions between bee species and floral morphology. However, dye transfer was more sensitive than pollen transfer to differences in floral morphology. CONCLUSIONS: The results from this study add to a growing body of literature highlighting that floral visitors vary in pollination effectiveness, and that visitors carrying the most pollen on their bodies may not always be the most efficient at depositing pollen on stigmas. Understanding the magnitude of variability in pollinator quality is one important factor for predicting how different pollinator taxa may influence the evolution of floral traits.     

Both morph‐ and species‐dependent asymmetries affect reproductive barriers between heterostylous species

The interaction between floral traits and reproductive isolation is crucial to explaining the extraordinary diversity of angiosperms. Heterostyly, a complex floral polymorphism that optimizes outcrossing, evolved repeatedly and has been shown to accelerate diversification in primroses, yet its potential influence on isolating mechanisms remains unexplored. Furthermore, the relative contribution of pre‐ versus postmating barriers to reproductive isolation is still debated. No experimental study has yet evaluated the possible effects of heterostyly on pre‐ and postmating reproductive mechanisms. We quantify multiple reproductive barriers between the heterostylous Primula elatior (oxlip) and P. vulgaris (primrose), which readily hybridize when co‐occurring, and test whether traits of heterostyly contribute to reproductive barriers in unique ways. We find that premating isolation is key for both species, while postmating isolation is considerable only for P. vulgaris; ecogeographic isolation is crucial for both species, while phenological, seed developmental, and hybrid sterility barriers are also important in P. vulgaris, implicating sympatrically higher gene flow into P. elatior. We document for the first time that, in addition to the aforementioned species‐dependent asymmetries, morph‐dependent asymmetries affect reproductive barriers between heterostylous species. Indeed, the interspecific decrease of reciprocity between high sexual organs of complementary floral morphs limits interspecific pollen transfer from anthers of short‐styled flowers to stigmas of long‐styled flowers, while higher reciprocity between low sexual organs favors introgression over isolation from anthers of long‐styled flowers to stigmas of short‐styled flowers. Finally, intramorph incompatibility persists across species boundaries, but is weakened in long‐styled flowers of P. elatior, opening a possible backdoor to gene flow through intramorph pollen transfer between species. Therefore, patterns of gene flow across species boundaries are likely affected by floral morph composition of adjacent populations. To summarize, our study highlights the general importance of premating isolation and newly illustrates that both morph‐ and species‐dependent asymmetries shape boundaries between heterostylous species.     

Is sexual organ reciprocity related to legitimate pollen deposition in distylous Pulmonaria (Boraginaceae)?

Heterostyly, i.e. the reciprocal positioning of anthers and stigmas, is a floral polymorphism that is thought to promote disassortative (i.e. between‐morph) pollination and to maintain genetic diversity within populations. Recent research, however, has shown several cases of heterostylous plant species in which the reciprocal positioning of the sexual organs varies, which may affect the likelihood of ‘legitimate’ pollination between compatible morphs, and hence morph fitness and ultimately the stability of this floral polymorphism. In this study, we investigated floral morphology and stigmatic pollen deposition patterns in nine Pulmonaria species (Boraginaceae) across Europe to test the hypothesis that sexual organ reciprocity is related to legitimate pollen deposition. We used a recently developed adaptive accuracy concept to assess the level of reciprocity and to relate inaccuracies in positioning of sexual organs to legitimate pollen deposition measured on stigmas in the field. The nine investigated Pulmonaria species showed substantial deviations from perfect reciprocity, with total inaccuracies varying between 2.9 and 20.3% relative to the squared mean of all anther and stigma heights in the population. In almost all cases, most of the inaccuracy was generated by the high‐level organs. Total pollen deposition was always higher on stigmas of long‐styled individuals, but stigmas of short‐styled individuals received significantly more legitimate pollen than stigmas of long‐styled individuals (71.23% and 38.75% respectively). We also found a weak but significant relationship between measures of inaccuracy and the level of disassortative pollination. Under these conditions of pollen flow, the distylous mating system is predicted to persist in only eleven (27%) of all sampled populations. Overall, these results indicate that deviations from perfect reciprocal herkogamy are common in Pulmonaria and have an impact on legitimate pollen deposition.     

Parental effects and gender specialization in a tropical heterostylous shrub

Abstract.— Male sterility in hermaphroditic species may represent the first step in the evolution toward dioecy. However, gender specialization will not proceed unless the male-sterile individuals compensate for fitness lost through the male function with an increase in fitness through the female function. In the distylous shrub Erythroxylum havanense , thrum plants are partially male-sterile. Using data collected throughout eight years, we investigated whether thrum individuals have an increased performance as female parents, thereby compensating for their loss of male fitness. We found that thrum plants outperformed pins in the probabilities of seed maturation and germination and long-term growth of the seedlings. In turn, pollen from pin plants achieved greater pollen tube growth rates. Our results suggest that the superior performance of the progeny of thrum maternal plants is a consequence of better seed provisioning via effects of the maternal environment, cytotype or nuclear genes. Overall, our results suggest that E. havanense is evolving toward a dioecious state through a gynodioecious intermediate stage. This evolutionary pathway is characterized by an unusual pattern of gender dimorphism with thrums becoming females and pins becoming males. We propose that this pattern may be better explained by the interaction between male-sterility cytoplasmic genes and the heterostyly supergene.     

Reproductive correlates of mating system variation in Eichhornia paniculata (Spreng.) Solms (Pontederiaceae)

Comparative studies of related plant species indicate that evolutionary shifts in mating systems are accompanied by changes in reproductive attributes such as flower size, floral morphology, and pollen/ovule ratio. Recent theoretical work suggests that patterns of investment in reproduction should also change with the mating system. In a glasshouse study, we investigated the extent to which mating system differences among populations of Eichhornia paniculata (Pontederiaceae) were correlated with changes in allocation to male and female function, floral display, and the regulation of investment in reproduction through fruit and ovule abortion. Significant differences in the amount of biomass allocated to reproductive structures were evident among six populations of E. paniculata. As predicted by sex allocation theory, the proportion of dry weight allocated to male function decreased with the outcrossing rate of populations. Six of the eight attributes used to characterize floral display also differed significantly among populations. However, with the exception of two attributes describing the number of flowers produced by inflorescences, these were not correlated with outcrossing rate. Levels of fruit and ovule abortion were determined in two populations with contrasting mating systems under different nutrient and pollination treatments. Virtually all fruits initiated by plants from a self-fertilizing population were matured, while the amount of fruit abortion in an outcrossing population increased with flower production. Ovule abortion was low in both populations. Our results demonstrate that the evolution of self-fertilization in E. paniculata is associated with changes in investment to reproduction that normally distinguish selfing and outcrossing species.     

Morph-biased male sterility in the tropical distylous shrub Erythroxylum havanense (Erythroxylaceae)

This study explores whether ecological factors, such as pollinators and pollen flow, or variation in pollen and ovule development account for the observed differences (approximately twofold) in the reproductive output of pin and thrum individuals of Erythroxylum havanense. The importance of ecological factors was assessed by means of comparison of the identity of pollinators and the rates of flower visitation, and by performing controlled hand pollinations and measurements of fruit set. In addition, we described the pollen and ovule development of thrum and pin individuals. Our results indicate that pollinators of E. havanense do not distinguish between floral morphs. The differences in fruit set between pin and thrum plants held even after hand pollination and, therefore, the observed differences in reproductive output between floral morphs of E. havanense cannot be explained in terms of asymmetrical pollen flow. There were no differences in the pattern of gynoecium development between the pin and thrum morphs, however androecium development showed marked differences between the morphs, and there was a resemblance between the developmental pathways leading to male sterility of the thrum morph of E. havanense with that of species with cytoplasmic male sterility (CMS).     

Quantitative evaluation of stigma polymorphism in a tristylous weed,Lythrum salicaria (Lythraceae)

Tristyly involves three different forms of flowers that differ reciprocally in the heights of stigmas and anthers within flowers. Apart from the style and stamen lengths, heterostylous species also demonstrate pollen and stigma polymorphisms. We quantified stigma polymorphism in tristylous Lythrum salicaria by measuring the stigma diameters, structure of papillae, and density and distribution of papillae on the stigma from flower samples of 201 individuals belonging to three morphs. The diameter of the stigma and the distribution of papillae were quantified using a scanning electron microscope, and the structure of papillae was determined using a light microscope. The stigma diameter in the long morph was significantly greater than in the mid and short morphs. While the density of stigmatic papillae was significantly greater in the mid and short morphs than in the long morph, the total number of papillae per stigma did not differ across morphs. The length and diameter of papillae at the apex, neck, and base were significantly greater in the long morph followed by the mid and short morphs. A discriminant function analysis separated the long morph from the mid and short morphs based on the canonical scores of measurements of papillae structure. The stigma polymorphism coupled with those of pollen may play a functional role in self-incompatibility mechanisms.