Trimorphic Populations Research Articles

The influence of floral morph ratios and low plant density on mating and fertility in a tristylous colonizing species

Sexual reproduction in heterostylous populations may be vulnerable to demographic conditions because of the small number of mating types in populations. Here, we investigate mating and fertility under natural and experimental conditions in tristylous Lythrum salicaria L., an invasive species that exhibits a wide range of floral morph ratios and demographic contexts. We grew 147 open-pollinated seed families from six populations with different morph structures to estimate intermorph mating (d). In a field experiment, we used progeny ratios from 47 spatially isolated individuals to estimate d, and measured the intensity of pollen limitation experienced by the morphs. The M- and S-morphs experienced high rates of d, regardless of population size or morph ratio. Estimates for the L-morph revealed low levels of intramorph mating in three dimorphic and two trimorphic populations, but near complete intramorph mating in a monomorphic population. Despite high levels of intermorph mating in the field experiment, the morphs experienced significant pollen limitation of fruit and seed set, but this did not differ in intensity among the morphs. Our field experiment demonstrates that although plant isolation was associated with pollen limitation of seed set, “long-distance” bee-mediated pollen flow served to maintain intermorph mating. Tristyly in L. salicaria is remarkably robust to the demographic variation associated with colonization.

Greater mycorrhizal colonization of unisexual morphs than of hermaphroditic morphs of Jacaratia mexicana during flowering and fruiting in central Mexico

Different sexual morphs of dimorphic and trimorphic populations of plants have different demands for resources correlating with phenotypic and ecological dissimilarities among morphs. Few studies have investigated the variations in mycorrhizal symbiosis across the sexual morphs of trimorphic plant populations. Here, we report seasonal variations in the mycorrhizal colonization of the roots of female, male and hermaphroditic trees of Jacaratia mexicana under field conditions in central Mexico. Mycorrhizal colonization was low in the rainy season, when J. mexicana bears only leaves, but high in the dry season when it flowers and fruits. In the dry season, the mycorrhizal colonization of female and male trees was over twofold higher than that of the hermaphroditic trees, and all three morphs had low mycorrhizal colonization in the rainy season. The lack of seasonal variation in the mycorrhizal colonization of hermaphroditic trees, together with other available evidence, supports the idea that hermaphroditic trees of J. mexicana might have some physiological disadvantages compared with male and female trees.

Reproductive strategy of the invasive Oxalis pes-caprae: distribution patterns of floral morphs, ploidy levels and sexual reproduction

Oxalis pes-caprae, a tristylous flowering plant native to South Africa, is described in the western Mediterranean basin as an asexual—only 5x short-styled morph (5x S-morph) invasive weed losing all mating partners after introduction. The objective of this study was to reassess the patterns of floral morph and cytotype distribution and the sexual reproduction ability in this invaded range. For that, floral morph and cytotype composition were evaluated in 39 populations of O. pes-caprae in a methodical sampling. The reproductive success of natural populations was assessed as fruit and seed production and seed germination for all floral morphs and cytotypes detected. Self- and morph-incompatibility were also studied with controlled hand pollinations. A remarkable diversity in floral morph and cytotype composition was observed. Furthermore, we observed successful sexual reproduction in several localities across the surveyed area. The S-morph is still dominant in this invaded area, and although it was mostly 5x, an additional cytotype (4x) was also recorded. Records of both a mid-styled morph (M-morph) and an area with trimorphic populations of this species are reported here for the first time in the invasive range of the Mediterranean basin. The long-styled morph appears to occur randomly across the surveyed area, while the M-morph is concentrated mainly in Estremadura province (Portugal), where a breakdown in the incompatibility system was observed. These distribution patterns may result from events of sexual reproduction after incompatibility breakdown and/or from multiple introduction events from the native area. The ability to reproduce sexually, undetected so far, may have important impacts in the population dynamics and major consequences for the adaptation and selection potential of O. pes-caprae in this invaded area.

Evolutionary pathways to self‐fertilization in a tristylous plant species

Evolutionary transitions from outcrossing to selfing occur commonly in heterostylous genera. The morphological polymorphisms that characterize heterostyly provide opportunities for different pathways for selfing to evolve. Here, we investigate the origins and pathways by which selfing has evolved in tristylous Eichhornia paniculata by providing new evidence based on morphology, DNA sequences and genetic analysis. The primary pathway from outcrossing to selfing involves the stochastic loss of the short-styled morph (S-morph) from trimorphic populations, followed by the spread of selfing variants of the mid-styled morph (M-morph). However, the discovery of selfing variants of the long-styled morph (L-morph) in Central America indicates a secondary pathway and distinct origin for selfing. Comparisons of multi-locus nucleotide sequences from 27 populations sampled from throughout the geographical range suggest multiple transitions to selfing. Genetic analysis of selfing variants of the L- and M-morphs demonstrates recessive control of the loss of herkogamy, although the number of factors appears to differ between the forms. Early stages in the establishment of selfing involve developmental instability in the formation of flowers capable of autonomous self-pollination. The relatively simple genetic control of herkogamy reduction and frequent colonizing episodes may often create demographic conditions favouring transitions to selfing in E. paniculata.

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.

Mating patterns and demography in the tristylous daffodil Narcissus triandrus

Mating patterns in plant populations are influenced by interactions between reproductive traits and ecological conditions, both factors that are likely to vary geographically. Narcissus triandrus, a wide-ranging heterostylous herb, exhibits populations with either two (dimorphic) or three (trimorphic) style morphs and displays substantial geographical variation in demographic attributes and floral morphology. Here, we investigate this variation to determine if demography, morphology, and mating system differ between the two sexual systems. Our surveys in Portugal and NW Spain indicated that dimorphic populations were less dense, of smaller size, and had larger plants and flowers compared to trimorphic populations. Outcrossing rates estimated using allozyme markers revealed similar outcrossing rates in dimorphic and trimorphic populations (t(m) dimorphic=0.759; t(m) trimorphic=0.710). All populations experienced significant inbreeding in progeny (mean F=0.143). In contrast, parental estimates of inbreeding were not significantly different from zero (mean F=0.062), implying that few inbred offspring survive to reproductive maturity due to inbreeding depression. Although the majority of inbreeding results from selfing, significant levels of biparental inbreeding were also detected in eight of the nine populations (mean s(s)-s(m)=0.081). Density was negatively associated with levels of selfing but positively associated with biparental inbreeding. Population size was positively associated with outcrossing but not biparental inbreeding. There were no consistent differences among the style morphs in outcrossing or biparental inbreeding indicating that the maintenance of trimorphism vs dimorphism is unlikely to be associated with inbreeding of maternal parents.

Severely reduced sexual reproduction in northern populations of a clonal plant, Decodonverticillatus (Lythraceae)

Summary In flowering plants the balance between sexual and clonal, asexual reproduction can vary widely. We quantified variation in sexual reproduction in a tristylous, clonal, aquatic plant, Decodon verticillatus, and investigated the role of ecological and genetic factors in causing this variation. We surveyed components of sexual fertility and vegetative growth in 28 populations distributed along a 500‐km latitudinal transect in New England, USA. Northerly populations tend to be monomorphic (M) for style length, and probably therefore have reduced sexual reproduction compared with southerly, trimorphic (T) populations. Compared with T populations (n = 10), M populations (n = 18) exhibited large reductions for all components of sexual reproduction, including flower production, pollen deposition, pollen tube growth, fertilization, fruit set and seeds per fruit. Seven M populations produced no seed at all, and the other 11 very little (mean = 24 vs. 1139 seeds per plant in trimorphic populations). Clonal propagation was also greatly reduced in M populations. A survey of three polymorphic allozyme loci detected only single, usually heterozygous, genotypes in 15 M populations, whereas all T populations were genotypically diverse. The other three M populations contained three or fewer genotypes and one always predominated. Sexual recruitment is therefore extremely rare. Comparison of the sexual fertility of M and T populations in a concurrent common glasshouse experiment with our field data revealed that reduced sexual performance in northern M populations is principally due to genetic factors, but is also caused by ecological factors that covary with latitude. This abrupt shift away from sexual reproduction in populations at the northern periphery of the geographical range in D. verticillatus may greatly limit their evolutionary potential and restrict further northward expansion.

LOSS OF SEX IN CLONAL POPULATIONS OF A FLOWERING PLANT, DECODON VERTICILLATUS (LYTHRACEAE).

The loss of traits that no longer increase fitness is a pervasive feature of evolution, although detailed studies of the genetic, developmental, and evolutionary factors involved are few. Most perennial plants practice both sexual and clonal reproduction, and it has been hypothesized that populations with little sexual recruitment may lose the capacity for sexual reproduction by fixing mutations that disable one or more of the many processes involved in sex. The clonal, tristylous aquatic plant, Decodon verticillatus, exhibits marked geographical variation in sexual recruitment. Populations at the northern limit of the range are usually monomorphic for style length consist of single genotypes, and produce almost no seed, due, in part, to environmental conditions that inhibit pollination, fertilization, and seed maturation. Controlled crosses in a greenhouse provided evidence for greatly reduced sexual capacity in an exclusively clonal, monomorphic population. Plants from this infertile population produced only 3-18% as many seeds per pollination as fertile populations. Observations of pollen tube growth indicated that infertility is due to severe reductions in pollen tube numbers both early after pollination and later when pollen tubes were traversing the ovary, due primarily to the inability of pistils to support normal tube growth. A three-year greenhouse experiment comparing fertility, survival, and growth of F1 progenies produced from reciprocal crosses between plants from the infertile population and those from nearby fertile populations suggested that the genetic basis for infertility is simple and may involve a single recessive mutation. In addition, the results did not reveal any association between infertility and other aspects of survival and vegetative vigor. The infertile genotype was likely fixed in the population through founder effect rather than indirect selection resulting from antagonistic pleiotropy or direct selection of advantages associated with reduced investment in sexual reproduction. A broader comparison of sexual fertility in 15 clonal, monomorphic populations and five genotypically diverse, trimorphic populations under greenhouse conditions revealed substantial infertility in all but one monomorphic population. Populations varied somewhat in the stage at which infertility was expressed, however, pollen tube growth was impaired in all populations. These results provide strong support for the hypothesis that complex traits like sex are degraded by mutation when they no longer increase fitness.

Snail arboreality: the microdistribution ofSitala jenynsi(Gastropoda: Ariophantidae)

The habitats and patterns of vertical migration of the shell banding morphs of the snail Sitala jenynsi (Pfeiffer) were studied in Dar es Salaam and Wazo regions of central coastal Tanzania Both dimorphic and trimorphic populations were arboreal throughout the year. The snails occurred randomly within mid-heights 180 to 360 cm. without apparent morph differences in micro-distribution. The pattern of circadial vertical distribution consisted of upward migrations during mornings and downward shifts during late evenings. These oscillations regressed negatively on relative humidity (RH), a unit advance in RH representing an eight-fold descent of snails on daytime height. Variations in RH accounted for much of the total variance in the mean vertical distance. During high RH, however, temperature and mating behaviour probably exerted major influences on snail micro-distribution. The effects of arboreality on snail survival and initiation of differentiating subpopulations are proposed.

Genetic control of fruit polymorphism in the GenusFedia (Valerianaceae) in the light of dimorphic and trimorphic populations ofF. pallescens

The genusFedia consists of three species (F. cornucopiae, F. graciliflora andF. pallescens) of winter annual herbs, endemic to the western Mediterranean Basin. The deciduous terminal fruits of these taxa are polymorphic in the development of their pericarp and/or calyx, and each population is dimorphic or more rarely trimorphic. The three main fruit types are dispersed in several manners, and are specialized for either epizoochory or myrmecochory. On the basis of our experimental study of dimorphic and trimorphic populations ofF. pallescens subsp.pallescens, a genetic model is presented in order to explain the control of this intrapopulational polymorphism. It is postulated that two diallelic loci are tightly linked on the same chromosome in a functional supergene. One allele of each locus displays a dominance effect in the heterozygous state. Within the four possible homologous allelic segments, only two are present in the dimorphic populations, three in the trimorphic ones, and are otherwise associated in diverse combinations in the remaining taxa of the genus. Similar examples of fruit polymorphism are already documented in the tribeValerianeae, subtribeFediinae. The hypothesis is put forward that this fruit polymorphism is a synapomorphy for the subtribe.

Evolutionary consequences of extensive morph loss in tristylous decodon verticillatus (Lythraceae): a shift from tristyly to distyly?

The evolution of distyly from tristyly has occurred repeatedly, especially in the Lythraceae. However, the evolutionary forces involved remain unclear since species exhibiting transitional stages between tristyly and distyly have rarely been studied. The self‐compatible, wetland perennial Decodon verticillatus (Lythraceae) may provide this transitional variation since populations commonly lack style morphs, particularly the mid‐styled (M) morph. In dimorphic populations lacking the M morph, anthers positioned at the mid level in both the long‐ (L) and short‐styled (S) morphs have lost their target stigma, setting the stage for either evolutionary repositioning of mid‐level anthers to increase pollen export to L and S stigmas, or increased variability in mid‐level anther position resulting from relaxed selection. We examined these two hypotheses by comparing floral morphology in eight dimorphic and ten trimorphic populations from throughout the species’ range. We found no evidence that loss of the M morph has led to evolutionary modification of mid‐level stamens. While mid‐level stamens of the S morph were 11.0 ± 4.0% (mean ± 1 SE) longer than those of the L morph in dimorphic populations, divergence in stamen length between morphs occurred to the same extent (10.4 ± 2.0%) in trimorphic populations and cannot be attributed to the absence of the M morph. Analyses of variability using median ratio tests revealed no difference in the variability of mid‐level stamen length between dimorphic and trimorphic populations. Mid‐level stamens were not more variable than long‐ and short‐level stamens within dimorphic populations. The consistent divergence in mid‐level stamens between the L and S morphs may reflect morph‐specific differences in the optimal position of mid‐level anthers for maximizing cross‐pollination and avoiding self‐fertilization.

POPULATION STRUCTURE AND MORPH-SPECIFIC FITNESS DIFFERENCES IN TRISTYLOUS LYTHRUM SALICARIA.

In tristylous plant populations, style-morph frequencies are governed by an interaction between frequency-dependent selection due to disassortative mating and stochastic processes. Provided that there are no inherent fitness differences among morphs, frequency-dependent selection should result in equal morph frequencies at equilibrium. Stochastic models indicate that the short-styled morph has the highest and the long-styled morph the lowest probability of being lost from local populations as a result of random processes. We surveyed the morph composition of 82 populations of the tristylous, self-incompatible herb Lythrum salicaria in two archipelagos, one in central and one in northern Sweden, located close to the range-margin of the species. To examine whether deviations from even morph frequencies can be explained by among-morph differences in reproductive success, we quantified flower and seed production in six and three populations in the northern and southern archipelago, respectively, and we recorded segregation ratios in offspring produced in six trimorphic populations in the northern area. Seed germination and offspring growth were studied in the greenhouse. Ninety percent of the populations in the southern archipelago (N = 31) and 69% of the populations in the northern archipelago (N = 35) were trimorphic; the remaining populations were dimorphic (only populations consisting of at least three flowering plants considered). Dimorphic populations were smaller than trimorphic populations, as predicted by stochastic models. There was a striking difference in the morph composition of L. salicaria populations between the two archipelagos. In the southern archipelago, there was a slight excess of the long-styled morph and a corresponding deficiency of the short-styled morph. In contrast, the northern populations were characterized by a marked deficiency of the mid-styled morph: the average frequency of the mid-styled morph in trimorphic populations was 0.21, and nine of eleven dimorphic populations lacked the mid-styled morph. In both archipelagos, the long-styled morph (the most common morph) produced about 20% fewer seeds per fruit than the other morphs. The long-styled morph also tended to produce fewer seeds per plant. A hand-pollination experiment performed in two of the northern populations indicated that seed production per flower was pollen-limited in the long-styled morph but not in the other two morphs. Seed germination and offspring size after 24 weeks of growth did not differ among morphs. The mid-styled morph tended to have a higher representation in the offspring than in the parental generation in all six trimorphic populations studied further indicating that the deficiency of the mid-styled morph in the northern archipelago does not represent an equilibrium. Taken together, the results do not support the hypothesis that morph-specific differences in reproductive success can account for deviations from even morph frequencies in L. salicaria. It is suggested that among-morph differences in other components of fitness and historical factors may contribute to the current morph structure.

Stamen elongation, pollen size, and siring ability in tristylous elchhornia paniculata (Pontederiaceae)

Selfing variants in tristylous Eichhornia paniculata (Pontederiaceae) possess an elongated, short‐level stamen adjacent to mid‐level stigmas, which causes autonomous selfing. The variants commonly spread in dimorphic, but not trimorphic populations in northeast Brazil. We investigated the effect of stamen elongation on pollen size and siring ability. Competition experiments using controlled hand‐pollination and allozyme markers were used to compare different pollen types. Pollen from the elongated stamen was significantly larger in size than pollen from unmodified short‐level stamens. In mixed pollinations of mid‐level stigmas, pollen from the elongated stamen sired significantly more seed than pollen from unmodified short‐level stamens. Despite these differences, the size and compatibility of pollen from the elongated stamen were more similar to short‐ than mid‐level pollen, indicating that alterations to stamen level were not associated with major changes in pollen characteristics. The results suggest that the advantage of selfing variants in dimorphic populations is mainly due to efficient pollen transfer to mid‐level stigmas rather than increased postpollination siring success of pollen from modified stamens. In addition, the absence of major changes in pollen size and compatibility associated with stamen elongation support other lines of evidence indicating that selfing variants are not the result of recombination in the putative heterostyly supergene.

Style Morph Ratios in Tristylous Decodon Verticillatus (Lythraceae): Selection vs. Historical Contingency

Tristylous plant populations should exhibit equal frequencies of the three style morphs at equilibrium. In contrast, New England and central Ontario populations of Decodon verticillatus (Lythraceae) show a marked deficiency of the mid—styled (M) morph. This pattern was also evident in an independent sample of 30 populations from eastern Ontario: all nine cases of morph loss involved the M morph, and this morph also occurred at low frequencies, especially in large trimorphic populations. The hypothesis that a fitness disadvantage to the M morph accounts for its low frequency was not supported by data from a series of field and glasshouse experiments. Measures of reproductive success from two populations showed no disadvantage to the M morph, but revealed substantially higher seed set in the mid— (M) and short—styled (S) morphs than in the long—styled (L) morph. A pollen addition experiment showed that low seed set in the L morph can only partly be explained by reduced pollen receipt. Comparison of morph frequencies between adult plants and their open—pollinated progeny in three populations failed to reveal any morph—specific fitness differences. Analysis of growth and flowering of open—pollinated progeny from three populations in a 2—yr glasshouse experiment revealed differences among morphs for some parameters in some populations, but no consistent patterns were evident. The deficiency of the M morph may have resulted from an historical accident during post—glacial colonization of parts of the northern range. Computer calculations examining the return of morph frequencies to equilibrium after major perturbations indicated that populations of long—lived clonal species like D. verticillatus may preserve skewed morph ratios for >10 000 yr. Moreover, the distribution of fossil seeds indicates that this species has occurred in its post—glacial range in southern Ontario for only °5000 yr. It is important to distinguish between ecological and evolutionary time scales when testing selective interpretations of microevolutionary patterns involving clonal taxa in glaciated regions.

POLLEN DISCOUNTING AND THE SPREAD OF A SELFING VARIANT IN TRISTYLOUS EICHHORNIA PANICULATA: EVIDENCE FROM EXPERIMENTAL POPULATIONS.

Floral traits that increase self-fertilization are expected to spread unless countered by the effects of inbreeding depression, pollen discounting (reduced outcross pollen success by individuals with increased rates of self-fertilization), or both. Few studies have attempted to measure pollen discounting because to do so requires estimating the male outcrossing success of plants that differ in selfing rate. In natural populations of tristylous Eichhornia paniculata, selfing variants of the mid-styled morph are usually absent from populations containing all three style morphs but often predominate in nontrimorphic populations. We used experimental garden populations of genetically marked plants to investigate whether the effects of population morph structure on relative gamete transmission by unmodified (M) and selfing variants (M') of the mid-styled morph could explain their observed distribution. Transmission through ovules and self and outcross pollen by plants of the M and M' morphs were compared under trimorphic, dimorphic (S morph absent), and monomorphic (L and S morphs absent) population structures. Neither population structure nor floral morphology affected female reproductive success, but both had strong effects on the relative transmission of male gametes. The frequency of self-fertilization in the M' morph was consistently higher than that of the M morph under all morph structures, and the frequency of self-fertilization by both morphs increased as morph diversity of experimental populations declined. In trimorphic populations, total transmission by the M and M' morphs did not differ. The small, nonsignificant increase in selfing by the M' relative to the M morph was balanced by decreased outcross siring success, particularly on the S morph. In populations lacking the S morph, male gamete transmission by the M' morph was approximately 1.5 times greater than that by the M morph because of both increased selfing and increased success through outcross pollen donation. Therefore, gamete transmission strongly favored the M' morph only in the absence of the S morph, a result consistent with the distribution of the M' morph in nature. This study indicates that floral traits that alter the selfing rate can have large and context-dependent influences on outcross pollen donation.

Multiple origins of self‐fertilization in tristylous Eichhornia paniculata (Pontederiaceae): Inferences from style morph and isozyme variation

AbstractPopulations of Eichhornia paniculata (Pontederiaceae) exhibit a wide range of mating systems, from predominant outcrossing to high levels of self‐fertilization. The origin of self‐fertilization in this tristylous species is associated with the loss of style‐length morphs from populations and the spread of self‐pollinating, floral variants. We examined geographic variation in style morph and allozyme frequencies to determine whether the loss of style morphs and transition to selfing could have multiple origins in E. paniculata. Surveys of floral variation in 167 populations from six states in northeastern Brazil revealed that at least one style morph was absent from 29.3%. Non‐trimorphic populations occurred in all states and ranged in frequency from 9% in Ceará to 68% in Alagoas. Selfing variants occurred in 8.5% and 55% of trimorphic and non‐trimorphic populations, respectively, and were distributed among five of six states with primary concentrations in Alagoas and Pernambuco. A comparison of electrophoretic variation at 24 isozyme loci in 28 trimorphic, 13 dimorphic and 3 monomorphic populations indicated that non‐trimorphic populations contained 84% of the allelic variation present in trimorphic populations and were markedly differentiated from one another. Analyses of genetic distance and the distribution of rare alleles indicated that non‐trimorphic populations were often more similar to neighbouring trimorphic populations than to one another. Populations with selfing variants occurred at low frequency in three genetically distinct parts of the range. These results, in combination with genetic and morphological evidence suggest that style morphs are lost repeatedly from populations of E. paniculata and that selfing variants may have originated on at least three separate occasions in northeastern Brazil.

EXPERIMENTAL STUDIES ON THE FUNCTIONAL SIGNIFICANCE OF HETEROSTYLY.

Heterostyly has been viewed as both an antiselfing device and a mechanism that increases the proficiency of pollen transfer between plants. We used experimental manipulation of the morph structure of garden populations of self-compatible, tristylous Eichhornia paniculata to investigate the function of floral polymorphism. Outcrossing rates (t), levels of intermorph mating (d), and morph-specific male and female reproductive success were compared in replicate trimorphic and monomorphic populations. In trimorphic populations, t and d averaged 0.81 (2 SE = 0.03) and 0.77 (2 SE = 0.03) respectively, with no difference in either parameter among morphs. Ninety-five percent of outcrossed seeds were therefore the result of intermorph fertilizations. Male reproductive success of the long-styled morph was low, especially in comparison with plants of the short-styled morph. Outcrossing rates for each morph were higher in trimorphic than monomorphic populations where t averaged 0.71 (2 SE = 0.01), 0.30 (2 SE = 0.04) and 0.43 (2 SE = 0.1) for the long-, mid-, and short-styled morphs, respectively. Seed set was lower in monomorphic populations, particularly those composed of the L morph, reflecting reduced pollen deposition. Floral polymorphism therefore increased both outcrossing rate and fecundity but the magnitude of the differences varied among morphs. If the ancestral condition in heterostylous groups resembled the L morph, as has been suggested, data from this study suggests that the selective basis for the establishment of floral polymorphism could have been increased pollen transfer rather than higher levels of outcrossing.

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

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

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

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

FACTORS INFLUENCING FREQUENCY OF THE MID-STYLED MORPH IN TRISTYLOUS POPULATIONS OF OXALIS ALPINA.

Genetic models and computer simulations suggest that modifications of incompatibility in tristylous populations of Oxalis alpina from southeastern Arizona should result in evolution of distyly, due to the inability of the mid floral morph to compete effectively with the short and long floral morphs. To evaluate this hypothesis, progeny testing of naturally pollinated individuals was carried out over a three-year period for populations with differing mid frequencies. In contrast to expectations, mids appeared in greater than expected proportions in many cases. Detailed analysis of results from two trimorphic populations where the mid form is relatively uncommon revealed that mid morphs produced far more mid progeny among their offspring than expected. Assuming that all compatible pollen has the same probability of deposition on appropriate stigmas, mid excesses were found among progeny of shorts and longs in these populations as well. In a third population where the mid form is abundant, mid excesses were not found among the progeny of mids, nor were mid excesses typical among the progeny of shorts and longs. Mid excesses among progeny of populations where the mid form is uncommon may explain retention of this form, despite incompatibility modifications that should favor evolution of distyly. Over-representation of mids may result from gametophytic selection favoring mid alleles. If this is the case, tristylous populations may be in equilibrium, with the frequencies of the three morphs determined by the balance of the opposing selective forces resulting from incompatibility modifications and gametophytic selection.