Tristylous Species Research Articles

Architectural constraints, male fertility variation and biased floral morph ratios in tristylous populations.

Tristyly is a genetic polymorphism in which populations are comprised of three floral morphs (mating types) differing reciprocally in sex-organ height. Intermorph (disassortative) mating governed by a trimorphic incompatibility system should result in 1:1:1 morph ratios at equilibrium, but both deterministic and stochastic processes can cause skewed morph ratios in tristylous populations. Here, we investigate mechanisms causing morph-ratio bias in Pontederia parviflora, an emergent aquatic native to tropical America. We compared reproductive traits among morphs and surveyed 71 populations to determine patterns of morph-ratio bias. We then used simulation models of morph-frequency dynamics to test the hypothesis that morph-specific differences in pollen production and their influence on male fertility can explain patterns of morph-ratio bias. Ninety-seven percent of populations that we sampled were tristylous, but with a significant excess of the short-styled morph and a deficiency of the long-styled morph. Atypically for a tristylous species, mid-level anthers of the short-styled morph produced over twice as much pollen compared with the corresponding anthers of the long-styled morph. Our computer models incorporating this difference in male fertility resulted in morph ratios not significantly different from the average frequencies from our survey suggesting that the short-styled morph is more successful than the long-styled morph in siring ovules of the mid-styled morph. We propose that the difference in male fertility between morphs may be a non-adaptive consequence of a developmental constraint caused by the architecture of tristyly in Pontederiaceae.

Floral morphs and seed production from hand-pollination in a population of Oxalis corymbosa in Taiwan

Oxalis corymbosa, a tristylous species, lacks seed-set in Taiwan. In a previous survey, six floral types (3 floral morphs in combination with two types of anthers) of 23 populations in Taiwan were assigned based on qualitative visual observation. In this study, we conducted experiments on one of the populations to pursue following objectives: to analyze the floral types of the population based on quantitative measurement and to test the hypothesis that the lack of seed-set is partially attributed to pollination limitation. The heights of anthers and stigmata were measured and analyzed. The number of viable pollen grains per anther was estimated. Pollen germination and seed-production after hand-pollination treatments were examined. Variations in anther and stigma heights were found in each type of previously assigned flowers. Unequal heights of reciprocal sexual organs between floral types were also found. Each level of anthers of MS-Y (mid-styled bearing yellow anthers) and SHS-Y (semi-homostylous bearing yellow anthers) flowers contained about 60 and 25 viable pollen grains, respectively; while anthers of SHS-W (semi-homostylous bearing white anthers) flowers had no pollen. Among hand-pollination treatments of inter-morph, cross-intra-morph and self-intra-morph crossings, only in one inter-morph crossing, i.e. SHS-Y (pollen receiver)×l/MS-Y [pollen donor, pollen from lower (l)-level anthers of MS-Y], pollen tubes were found entering micropyles of the ovules, and seeds were built. Thus, this species can produce seeds by hand-pollination. The result supports that limitation of pollen transfer at least partially contributes to the absence of seed-production in field populations of O. corymbosa in Taiwan.

Invasion Fosters Change: Independent Evolutionary Shifts in Reproductive Traits after Oxalis pes-caprae L. Introduction.

Biological invasions offer optimal scenarios to study evolutionary changes under contemporary timescales. After long-distance dispersal, exotic species have to cope with strong mate limitation, and shifts toward uniparental reproduction have been hypothesized to be selectively advantageous. Oxalis pes-caprae is a clonal tristylous species native to South Africa, and invasive in Mediterranean regions worldwide. It reproduces sexually and asexually but the importance of each strategy differs between ranges. Native populations reproduce mostly sexually while in invasive ones asexual reproduction is the prevailing strategy due to the dominance of pentaploid monomorphic populations. Nevertheless, two contrasting scenarios have been observed after introduction: transition toward clonality, and re-acquisition of sexuality fueled by multiple introductions of compatible mates. Here, we aimed to assess evolutionary changes of reproductive traits in O. pes-caprae invasive populations and evaluate whether these traits could be related with invasion success and prevalence of certain forms in the western Mediterranean basin. Sexual and asexual reproduction traits were quantified under optimal conditions in a common garden experiment including native and invasive sexual, predominately asexual, and obligated asexual individuals. Different reproductive, ecological, and genetic constraints created by long-distance dispersal seem to have generated different selective pressures in sexual and asexual traits, with our results supporting evolutionary changes in invasive populations of O. pes-caprae. Native plants had higher sexual fitness, while a transition toward clonality was clear for invasive forms, supporting clonal reproduction as a major trait driving invasion. Differences were also observed among invasive plants, with sexual forms having increased dispersal potential; thus, they are expected to be in advantage in comparison with predominantly asexual and obligated asexual plants, and may become widespread in the future. Historical processes, like the initial introduction of predominantly asexual forms followed by sexual forms more recently, could be in the origin of current distribution patterns of O. pes-caprae in the western Mediterranean. This study shows that invasion processes are very dynamic and that ecological and genetic constraints determined by the invasion process may originate different reproductive strategies that are likely to determine invasion success.

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.

Foraging Behavior of Apis Mellifera (Hymenoptera: Apidae) and Lycastrirhyncha Nitens (Diptera: Syrphidae) onPontederia sagittata(Commelinales: Pontederiaceae) on a Disturbed Site

By influencing the exchange of pollen, floral visitor behavior largely promotes the reproductive success of the plants. Pontederia sagittata (C. Presl) (Commelinales: Pontederiaceae) is a tristylous species whose morphs (long-styled L, mid-styled M and short-styled S) differ in the arrangement of reproductive organs and the amounts and accessibility for food for pollinating insects. We evaluated the behavior of 2 common contemporary visitors to inflorescences, the exotic bee Apis mellifera (L.) (Apidae), a not historical pollinator, and the flower fly Lycastrirhyncha nitens (Bigot) (Syrphidae), a frequent visitor already reported on this aquatic plant, based on HD video records of the number of individuals and the frequency and duration of their visits to 300 inflorescences. Both species of insects preferred to visit S-morph inflorescences. Pollen collection and nectar feeding were the most important activities of the bees, whereas flower flies were observed fed only on nectar. Thus, these behaviors could play an important role in reproduction in the study population of P. sagittata.

Sexual reproduction of the pentaploid, short‐styled Oxalis pes‐caprae allows the production of viable offspring

Reproduction is a key factor for the successful establishment and spread of introduced species. Oxalis pes-caprae is a tristylous species with a self- and morph-incompatibility sexual system that, in the invaded range of the western Mediterranean Basin, has been found to reproduce asexually because only the pentaploid, short-styled morph (5x S-morph) was introduced. The objective of this study was to test the ability of the 5x S-morph of O.pes-caprae to produce viable offspring in the absence of compatible mates, exploring the hypothesis that new morphs could have emerged by sexual reproduction events of the initially introduced morph. Pollen germination, pollen tube development, fruit and seed production, seed germination and offspring ploidy levels were analysed after controlled hand-pollinations to assess self- and morph-incompatibility and production of viable gametes by the 5x S-morph. The self-incompatibility system is still operating, but a partial breakdown in the morph-incompatibility system combined with the production of viable gametes was observed, allowing sexual reproduction of the 5x S-morph in the invaded range. The ability of the 5x S-morph to reproduce sexually may have major consequences for the dynamics of invasive populations of O.pes-caprae and could be one of the factors involved in the occurrence of new floral morphs in this invaded range.

Pollination of invasive Eichhornia crassipes (Pontederiaceae) by the introduced honeybee (Apis mellifera L.) in South China

Heterostyly functions as an outcrossing mechanism facilitating accurate pollen transfer from anthers to stigmas of particular heights as a result of the behavior of specialist pollinators. However, heterostylous plants are also visited by generalist pollinators, which may affect the plant–pollinator mutualism. Eichhornia crassipes is a tristylous invasive species, with only the mid- and long-styled morphs (M and L) found in China. We recorded flower-visiting insects in Zhuhai, Zhongshan and Nanning in South China. We hand-pollinated the two morphs to determine their compatibility. In addition, by allowing controlled insect pollination in artificial isoplethically monomorphic and bimorphic populations, we undertook a detailed analysis of pollen deposition between the floral morphs, and fruit and seed set. Ranked by relative abundance, the flower-visiting insects were: Apis mellifera, A. cerana, Lasioglossum sp. and Eristalis arvorum. Hand pollination showed that both the M and L morphs were self-compatible, but the former was probably more so than the latter. Intra-morph pollen transfer by A. mellifera within a population was significantly greater than legitimate pollen transfer between populations, suggesting that the pollen exchange between populations was limited. Seed set of the L morph was significantly greater than that of the M morph in monomorphic populations, indicating intra-morph pollen deposition in the former was higher than in the latter. The results showed that A. mellifera was the major pollinator in South China and able to pollinate E. crassipes legitimately and to promote its fruit and seed set, even though high levels of intra-morph pollination occurred.

Trimorphic Incompatibility inPontederia subovata(Pontederiaceae): An Aquatic Macrophyte from Lowland South America

Premise of research. Tristyly is a floral polymorphism that is reliably reported from six flowering plant families. Populations of tristylous species often contain three floral morphs that differ reciprocally in stigma and anther height. Trimorphic incompatibility and a suite of ancillary pollen and stigma polymorphisms are commonly associated with sex organ trimorphism. Here, we investigate the tristylous syndrome of Pontederia subovata (Pontederiaceae), a little-known aquatic from southwest Brazil.Methodology. We conducted measurements of sex organ position, pollen size, pollen production, and stigmatic papillae length on the three floral morphs under glasshouse conditions and carried out a controlled pollination program to investigate whether P. subovata possesses a trimorphic incompatibility system and whether its expression varies among the floral morphs.Pivotal results. Pontederia subovata displays reciprocal stigma and anther heights and trimorphism in the size of pollen grains and the length of stigmatic papillae. Controlled pollinations provided evidence of trimorphic incompatibility: self- and intramorph crosses yielded significantly less seed than do intermorph crosses when pollen from anthers and stigmas of equivalent height were employed. However, trimorphic incompatibility was weakly expressed in the mid-styled morph when pollinations involved long-level anthers. The occurrence of stem-borne tubers is reported for the first time from Pontederiaceae.Conclusions. Our results confirm the presence of tristyly in P. subovata. The species has several features in common with other tristylous members of Pontederia, including striking variation in the strength of incompatibility among the floral morphs. We suggest that weak incompatibility in the mid-styled morph may provide reproductive assurance during colonization. Tuber formation in P. subovata probably evolved as an adaptation enabling persistence of genets during the marked dry season that characterizes the aquatic habitats in which the species occurs.

Variation in floral morph ratios in tristylous Oxalis squamata (Oxalidaceae): an Andean alpine endemic

Negative frequency-dependent selection resulting from disassortative mating should result in equal morph ratios in tristylous populations at equilibrium. However, surveys of morph-frequency variation in tristylous species commonly report deviations from equality. Here, we report variation in morph ratios in Oxalis squamata Zucc., an endemic of the Andean region of Chile and Argentina. Absence of clonal propagation in this species allows unambiguous estimates of the morph ratio of genets. We sampled floral morph ratios in 20 populations occurring in Central Chile and investigated the relation between morph evenness in populations and their size. All populations of O. squamata were tristylous but with significant heterogeneity among populations in morph ratios. Although small populations exhibited a greater variance in morph evenness, biased ratios were also evident in several large populations. We found no evidence of morph loss or a consistent bias in morph frequencies as reported in some tristylous species. Biased morph frequencies in large populations probably arise from episodic sexual recruitment following disturbance and a slow approach to equilibrium.

Pollen and ovule production, floral nectary structure, and nectar secretion dynamics in tristylous Lythrum salicaria L.

Lythrum salicaria L. (Lythraceae) is tristylous, each plant forming one of three floral morphs that differ in reciprocal placement of the stigma and two sets of anthers. Several reproductive traits were compared quantitatively among these morphs. Although mean pollen viability (≥93%) and total pollen per mature, indehiscent anther within a staminal level (mean CV = 11%) were constant, the patterns of mean pollen production per anther were complex, being significantly lowest in long stamens (1,490 grains) of the short-styled morph, but highest in intermediate stamens (3,590) of the long-styled morph. Overall, pollen production was greatest (38,200) in long-styled flowers and least (22,000) in short-styled ones. On the contrary, ovule quantities per ovary (mean 107) were similar among floral morphs; thus, pollen-to-ovule ratios spanned 192 (short-styled morph) to 364 (long-styled morph), relatively low values for a strictly xenogamous species. Each morph had a recessed annular nectary of similar dimensions encircling the ovary base, with equal numbers of modified stomata distributed uniformly on the nectary surface. Most stomata were solitary (94%), whereas 5% occurred in pairs and 1% of stomatal units had just one guard cell. During nectar secretion, about 16% of pores were closed plus 28% of pores were fully occluded. Similarly, nectar volumes and solute concentrations, peak rates of nectar secretion (mean 72–79 μg sugar h−1) at early afternoon, and the nectar’s sucrose prevalence [S/(G + F) ≥4.3] were not significantly different among morphs. Based on these similarities in nectary structure and nectar-secretion dynamics, traits rarely studied in tristylous species, the preferential visitation of any particular floral morph of L. salicaria during nectar foraging by insects is unlikely. Indeed, lack of discrimination among morphs by potential pollinators may be a key tenet of successful sexual reproduction in tristylous species.

Genetic Control of Floral Morph in Tristylous Pickerelweed (Pontederia cordata L.)

Pickerelweed (Pontederia cordata L.) is a diploid (2n = 2x = 16) tristylous aquatic perennial. Populations usually contain 3 floral morphs that differ reciprocally in style length and anther height (referred to as the long-, mid-, and short-styled morphs, hereafter L-, M-, and S-morphs). The floral polymorphism promotes disassortative mating among the 3 floral morphs and is maintained in populations by negative frequency-dependent selection. The objective of this study was to determine the number of loci, number of alleles, and gene action controlling floral morph in pickerelweed. Three parental lines (one each of the L-, M-, and S-morph) were used to create S1 and F1 populations. F2 populations were produced through self-pollination of F1 plants. Progeny ratios of S1, F1, and F2 generations revealed that tristyly is controlled by 2 diallelic loci (S and M) with dominant gene action. The S locus is epistatic to the M locus, with the S-morph produced by plants with the dominant S allele (genotype S _ _ _). Plants with recessive alleles at the S locus were either L-morph (ssmm) or M-morph (ssM_). The results of this experiment demonstrate that the inheritance of tristyly in pickerelweed is the same as previously reported for several tristylous species in the Lythraceae and Oxalidaceae.

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.

Geographic variation in floral morphology and style‐morph ratios in a sexually polymorphic daffodil

Narcissus triandrus is a tristylous daffodil from the Iberian Peninsula that has striking geographical variation in floral morphology and style-morph ratios. Here, we investigate the relation between this variation and ecological factors to understand mechanisms governing morph ratios. We estimated morph ratios in 124 populations throughout the range of N. triandrus and measured 13 morphological traits in 35-78 populations. Sampling of morph ratios conducted over 2-10-yr intervals demonstrated strong temporal stability. Variation in floral and vegetative traits enabled statistical prediction of morph-frequency variation among populations. Latitudinal gradients in precipitation and temperature were correlated with plant and flower size, with larger flowers in northern populations associated with bumblebee visitation and stylar dimorphism. Flowers of the L- and M-morphs differed significantly from the S-morph in several size-related characters, unlike other tristylous species. This pattern and the similarity in anther positions of the L- and M-morphs suggest that the M-morph of N. triandrus originated through genetic modifiers that shortened styles of the L-morph. Our findings support the hypothesis that geographical variation in style-morph ratios is largely governed by climatic gradients in the Iberian Peninsula, which influence the floral morphology and pollination biology of N. triandrus populations.

The evolution of distyly from tristyly in populations of Oxalis alpina (Oxalidaceae) in the Sky Islands of the Sonoran Desert

The evolution of distyly from tristyly was investigated in populations of Oxalis alpina at high elevations throughout the Sky Islands of the Sonoran Desert. Incompatibility systems in tristylous populations, where self-incompatible short-, mid-, and long-styled morphs occur in populations, vary from those typical of tristylous species in which each morph is equally capable of fertilizing ovules of the other two morphs, to breeding systems in which incompatibility relationships are asymmetric. In these populations, selection against the allele controlling expression of the mid-styled morph is likely. The degree of modification of incompatibility in the short- and long-styled morphs in 10 populations was strongly associated with fewer mid-styled morphs, supporting models predicting the effect of these modifications of incompatibility on frequency of the mid-styled morph. Self-compatibility of the mid-styled morph may be important for maintaining the frequency of this morph, depending on the level of self-pollination, self-fertilization, and the extent of inbreeding depression. Modifications of incompatibility in tristylous populations and the distribution of distylous populations of O. alpina in the Sky Island region have similar geographic components, indicating the potential importance of historical factors in the evolution of distyly from tristyly.

Oxalis debilis in China: Distribution of Flower Morphs, Sterile Pollen and Polyploidy

Oxalis debilis is a South American tristylous species that is currently naturalizing in China. Based on field observations and herbarium data a report is made on its pollination, morph frequencies, pollen viability, seed set and chromosome number. In addition, a new chromosome count for the species in Africa is provided. Field observations were conducted in six provinces in south-east China. Flower visitors were recorded and nectar sugar was measured with a refractometer. The species' compatibility system was determined by carrying out experimental self- and cross-pollinations on bagged inflorescences. Stigma receptivity and pollen viability was determined using the MTT test. Populations of O. debilis in China contain either the mid-style-length morph or the short morph, but not both. Pollination is by nectar- and pollen-foraging bees; pollen viability is low; and seed set in natural and experimentally self- or cross-pollinated flowers is extremely low. Chromosome counts indicate that O. debilis contains diploid and tetraploid forms in its native as well as introduced range, which does not support a previous hypothesis that the predominant vegetative reproduction in this species is an escape from pentaploidy.

CORRELATED EVOLUTION OF FLORAL MORPHOLOGY AND MATING-TYPE FREQUENCIES IN A SEXUALLY POLYMORPHIC PLANT

In sexually polymorphic species, reproductive morphology governs mating patterns and the character of negative frequency-dependent selection. If local environmental conditions cause sexual morphs to differ between populations, then frequency-dependent selection should create corresponding geographic variation in morph frequencies. We investigate this relation with a model of morph-ratio evolution and analysis of geographic variation in the heterostylous plant Narcissus triandrus. Unlike other tristylous species, N. triandrus possesses both imperfect reciprocity among morphs in sex-organ position and a self-incompatibility system that permits outcrossing within and between morphs. We sampled 137 populations throughout the Iberian Peninsula for floral-morph ratios, and measured floral morphology in 31 populations. Morph ratios exhibited three atypical features: (1) predominance of the long-styled (L) morph; (2) absence of the mid-styled (M) morph from 17.5% of populations; and (3) a negative relation between the frequencies of the L and M morphs among populations. Morph ratios varied geographically, with decreasing frequency of the M morph from the southeast to the northwest of the species' range. Much of this variation accompanied allometric change in the positions of sex organs, especially the mid-level organs, with the M morph declining in frequency and ultimately being lost in large-flowered populations. Using multivariate multiple regression, we demonstrate that variation in floral morphology among populations predicts this geographic variation in morph frequencies. Our theoretical analysis illustrates that patterns of pollen transfer governed by imperfect sex-organ reciprocity can select for unequal equilibrium morph ratios like those observed for N. triandrus. We interpret the L-biased morph ratios and the unusual morphology of N. triandrus as a consequence of its atypical intramorph compatibility system.

Trimorphic incompatibility in Eichhornia azurea (Pontederiaceae)

Eichhornia azurea (Pontederiaceae) is a mat- forming, clonal aquatic that inhabits lakes, marshes and river systems in many parts of the Neotropics. The spe- cies is tristylous with long-, mid-, and short-styled morphs commonly represented in natural populations. To investigate whether E. azurea possesses a trimorphic in- compatibility system typical of tristylous species, we conducted a controlled pollination experiment on 15 clones representing the three style morphs from a natural population near Rosario, Argentina. Comparisons of fruit and seed set following self-, illegitimate, and legitimate pollinations clearly demonstrated the presence of trimor- phic incompatibility in E. azurea. Self- and illegitimate pollinations produced significantly less fruit and seed than legitimate pollinations in all three style morphs. Pollen from the two anther levels within a flower exhib- ited contrasting compatibility relations in self-pollina- tions. In common with several other tristylous species in Pontederiaceae, the expression of self-incompatibility was weakest in the mid-styled morph and strongest in the short-styled morph. We discuss the ecological and evolutionary significance of the partial expression of trimorphic incompatibility in E. azurea.

Legitimate pollination of the tristylous flowers ofEichhornia azurea (Pontederiaceae) byAncyloscelis gigas bees (Anthophoridae, Apoidea)

Eichhornia azurea is a tristylous species of Pontederiaceae. Moderate self-incompatibility accompanies the floral heteromorphism of this species.Eichhornia azurea is almost exclusively visited by one bee species,Ancyloscelis gigas (Anthophoridae), in South Brazil. This species has an extremely long proboscis covered with recurved hairs. Bees use these hairs to collect pollen from the anthers placed inside the narrow perianth tube ofE. azurea. Analyses of the pollen load ofA. gigas females show that they are flower constant and carry pollen from all three anther levels. The behavior ofA. gigas and its proboscis morphology make this bee species narrowly adapted and an effective pollinator for facilitating legitimate pollination ofE. azurea flowers in southern Brazil.

Developmental aspects of tristyly in Lythrum salicaria

The developmental basis of floral polymorphism was investigated in the tristylous invasive species, Lythrum salicaria L. (Lythraceae). In tristylous species, the stigmas are positioned above (in the long morph), below (in the short morph), or between (in the mid morph) the outer and inner staminal whorls. Flower samples were collected at three different growth stages (early, pre-anthesis, and post-anthesis) from three genotypes from each of the three morphs to observe morph-specific differences in growth patterns of filaments and styles and their constituent epidermal cells. From each flower, I measured the length of styles and two types of stamens and their epidermal cells at the basal, intermediate, and apical regions of each organ. Differentiation of organ levels begins at a very early stage. Growth rate of the long pistil is higher than in the mid pistil followed by the short pistil. However, the growth rate of epidermal cells is higher in the short style followed by the mid and long styles. The number of cells does not increase during style development in the short morph but does increase in the long and mid morphs. Although the relative growth of the outer stamens is greater than in the inner stamens in all three morphs, the relative cell size is greater in the inner stamens than in the outer stamens. Cell size differs between outer and inner stamens in the long and mid morphs but not in the short morph. The intermediate cells are larger compared with the basal and apical cells of the stamens and styles. The number of epidermal cells increases in the outer stamens during development, whereas it remains constant in the inner stamens of the mid morph and increases only slightly in the inner stamens of long morph.Key words: floral development, growth and division of epidermal cells, heterostyly, purple loosestrife, style and stamen growth, style-stamen polymorphism.

Tristyly in the endangered Mascarene Island endemic Hugonia serrata (Linaceae)

Tristyly is a rare floral polymorphism known to occur in only five flowering plant families. One unresolved and potential additional case of tristyly concerns the genus Hugonia in the Linaceae. Here we confirm the existence of tristyly in the genus by reporting floral measurements made on Hugonia serrata Lam., an extremely rare species endemic to the Mascarene Islands of La Réunion and Mauritius in the southern Indian Ocean. We conducted an extensive search of all natural habitats on La Réunion Island where the species had been previously reported. Twenty‐eight individuals were found, of which nine were in flower. Of the nine flowering individuals five had long‐styled flowers with the stigmas placed above the two levels of anthers, three had intermediate length styles with the stigmas placed between the two anther levels, and one had stigmas placed below the two anther levels, i.e., three floral morphs could be identified based on the sequence of stigma and anther positions. Reciprocity and precision indices calculated for sexual organ length in each morph confirm that this variation is well within the range of values observed by previous workers on other tristylous species in other families. Our empirical data confirm the existence of tristyly in the genus Hugonia, thereby raising the number of known families in which tristyly occurs to six. Pollen size, pollen number, and anther length increased slightly with stamen length, but the low number of plants precludes statistical tests of these trends. A bibliographic survey suggests that tristyly may occur in several other species of the genus.