Interaction between founder effect and selection during biological invasion in an aquatic plant

Long-distance colonization and rapid range expansion associated with biological invasion may have major evolutionary consequences via both stochastic processes and selection. Using large-scale population genetic surveys, we demonstrate a major shift in the relative frequency of sexually fertile diploid versus sexually sterile triploid populations associated with the invasion of North America by a clonal aquatic plant, Butomus umbellatus. Most populations across the native European range were triploid (84% of 108), whereas most introduced populations were diploid (71% of 136). We evaluated the roles of stochastic processes versus natural selection in causing this shift by surveying predominantly neutral genetic variation at 28 RAPD loci. In Europe (EU) we detected 47 distinct genotypes among 142 plants sampled from 71 populations, whereas in North America (NA) we detected only six genotypes among 138 plants from 69 populations. Of the six NA genotypes, a set of four closely related genotypes were found only in triploid populations and a pair of closely related genotypes were found only in diploid populations, and these were genetically divergent from the triploid genotypes. This result is consistent with severe founder effect. Because sex creates genotypic variation and produces offspring with greater dispersal potential than those produced clonally, we tested the hypothesis that sexual reproduction characteristic of diploids has given them a colonization advantage that accounts for their high frequency in NA. However, we found little or no evidence of sexual recruitment in introduced diploids. One very widespread heterozygous genotype occurred in 95% of 38 introduced diploid populations (i.e., 72 of 76 plants surveyed) suggesting predominant clonal reproduction. Moreover genotypic diversity was not higher within or among diploid than triploid populations in either the native or introduced range. Low genetic diversity in diploid populations was also supported by a comparison of within-population quantitative variation for plant size under a common greenhouse environment. Thus, diploids have not been favored during colonization owing to their sexual fertility. However, concurrent studies have shown that NA diploids exhibit a much higher capacity for clonal reproduction, via small vegetative bulbils, than NA triploids, which almost never produce bulbils. The same difference in clonal capacity is not a consistent feature of the native EU populations. Taken together, these results suggest that strong founder effect has set the stage for a major increase in diploid frequency due to the particular, and possibly idiosyncratic, features of the diploid and triploid lineages introduced to North America.     

Causes and consequences of extreme variation in reproductive strategy and vegetative growth among invasive populations of a clonal aquatic plant, <Emphasis Type="Italic">Butomus umbellatus</Emphasis> L. (Butomaceae)

The mode of reproduction may influence the spread of invasive species by affecting evolutionary potential and dispersal ability. We sampled 51 introduced North American populations of the clonal aquatic plant Butomus umbellatus L. (flowering rush) and found extreme variation in sexual fertility caused by polyploidy. Populations consisted of diploids that produced thousands of viable seeds or of sexually sterile triploids. Although a trade-off between sexual and clonal reproduction predicts that the sexual sterility of triploids would be compensated for by greater clonal reproduction, a greenhouse experiment involving eight diploid and 10 triploid populations showed that diploid plants not only invest substantially in sexual structures but also make hundreds of tiny clonal bulbils on both rhizomes and inflorescences. In contrast, triploids do not make bulbils and have very limited scope for clonal multiplication and dispersal. Diploid populations were more frequent than triploid populations, especially in the Great Lakes region. This is probably because of the difference between cytotypes in clonal rather than sexual reproduction, as genetic analyses indicate a general lack of sexual recruitment in North America. Although triploids were less common, they have a wider geographical distribution. This could be due to a greater ecological tolerance resulting from polyploidy. However, genetic evidence suggests that triploids have become widespread via their use in and escape from horticulture. North America is being colonized by two distinct forms of B. umbellatus that differ strongly in reproductive strategy as well as the vectors and pathways of invasion.     

Isozyme variation under different modes of reproduction in two clonal winter annuals, Sedum rosulato-bulbosum and Sedum bulbiferum (Crassulaceae)

We studied isozyme variation in two annual species that produce bulbils, Sedum rosulato-bulbosum , which includes both sexually reproducing plants and obligate clonal plants that result from triploidy (fertile and sterile S. rosulato-bulbosum , respectively), and an obligate clonal plant, Sedum bulbiferum , to examine the relationship between reproductive mode and isozyme variation. The sterile S. rosulato-bulbosum population exhibited no genotypic variation, but showed high genetic variation (gene diversity, H _e  = 0.60) because five of the six loci that we analyzed were heterozygous. Almost all ramets of S. bulbiferum across 20 populations shared an identical isozyme phenotype, although we could not identify the genetic basis of the phenotype. In contrast, fertile S. rosulato-bulbosum exhibited genotypic variation across the species, but comprised genotypically uniform and polymorphic populations whose genotypic variations correlated positively with the genetic variations within the populations ( H _e at the genet level per population ranged from 0.08 to 0.37). Genetic drift and habitat conditions inhibiting seedling recruitment may have caused this among-population variation. The results for sterile and fertile S. rosulato-bulbosum suggest that exclusive clonal reproduction causes low genotypic variation, but maintains genetic variation within individuals. Factors that affect the maintenance of genetic variation in these plants are discussed on the basis of these findings.     

Evolutionary vestigialization of sex in a clonal plant: selection versus neutral mutation in geographically peripheral populations

The loss of traits that no longer contribute to fitness is widespread; however, the causative evolutionary mechanisms are poorly understood. Vestigialization could proceed through the fixation of selectively neutral degenerative mutations via genetic drift. Alternatively, selection may facilitate vestigialization if trait loss results in enhanced fitness. We tested these hypotheses using Decodon verticillatus, a clonal plant in which sexual sterility has arisen repeatedly in populations across the northern geographical range limit. We compared growth and survival of replicated genotypes from 7 sexually fertile and 18 sterile populations, over 3 years in a common environment. Survival of sterile genotypes was 53% greater than for fertile genotypes, but there was no difference in biomass accumulation. Almost all mortality, and hence increased performance of sterile genotypes, occurred during simulated overwinter dormancy. These observations suggest that selection has facilitated the vestigialization of sex, and thus do not support the neutral mutation hypothesis. The selective mechanism probably involves the relaxation of a genetic trade-off between sexual reproduction and survival: alleles that increase vegetative performance at the expense of sexual fertility are selected in geographically peripheral populations where sexual reproduction is suppressed by adverse environmental conditions.     

The importance of sexual and asexual reproduction in the recent evolution of Allium vineale

In the weedy plant species Allium vineale (wild garlic), individuals may simultaneously produce sexually and asexually derived offspring, by seed and bulbils, respectively. In this study, genetic and genotypic diversity was determined in samples from 14 European A. vineale populations using nuclear (RAPD) and cytoplasmic (PCR-RFLP of cpDNA) markers to investigate the importance of the different reproductive modes. In the whole sample, 77 nuclear multilocus genotypes and four chloroplast haplotypes (chlorotypes) were found. Populations exhibited a high degree of subdivision for nuclear and cytoplasmic markers as estimated from hierarchical F-statistics; at the same time, identical chlorotypes could be found in populations separated by large distances. Genotypic diversity was significantly lower than expected under free recombination in almost all populations, indicating that recruitment into populations is mostly by asexually produced offspring. Nevertheless, within each chlorotype, the distribution of markers from pairs of nuclear loci was incompatible with a purely clonal structure, suggesting that many multilocus genotypes have originated by sexual recombination rather than by mutation within asexual lineages. It is argued that the weedy habit of A. vineale is likely to have favored bulbil reproduction, whereas sexually generated genotypes may have facilitated local adaptation during the species' expansion across Europe.     

Clonality and recombination in the arctic plant Saxifraga cernua

The circumarctic clonal plant Saxifraga cernua reproduces efficiently via bulbils, largely depends on insects for pollination and appears to set seed very rarely. However, high levels of genotypic variation observed at small spatial scales in the arctic archipelago of Svalbard have been taken as evidence of occasional sexual reproduction. Here we assess the relative contributions of mutation and recombination to random amplified polymorphic DNA variation in four populations in East Greenland and re-analyse the Svalbard data. Greater variation due to recombination was predicted in Greenland than in Svalbard, because the higher summer temperatures and longer growing season likely increase the chances for sexual reproduction. Although we observed higher levels of genotypic diversity in Greenland than in Svalbard, matrix incompatibility and linkage disequilibrium measures provided no evidence of more sexuality, suggesting differences in glacial/postglacial history. The genetic structure and spatial distribution of clones suggest that clonal migration may increase variability in local populations, which is consistent with frequent large-scale migration in this species inferred from a circumarctic analysis of chloroplast DNA haplotypes. We conclude that a combination of somatic mutations and sexual reproduction has contributed to the observed patterns of genotypic diversity in the Greenland and Svalbard populations of S. cernua , and that sexual reproduction is important in maintaining genotypic diversity, despite the rarity of observations of seed setting.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 209–217.     

Evolutionary increase in sexual and clonal reproductive capacity during biological invasion in an aquatic plant Butomus umbellatus (Butomaceae)

To test the hypothesis that increased allocation to reproduction is selected during biological invasion, we compared germination, survival, growth, and reproduction of native vs. introduced populations of the invasive aquatic plant Butomus umbellatus in a common greenhouse environment. Although seedling emergence and establishment did not differ consistently, survival thereafter was twice as high for eight introduced North American than eight native European populations. As predicted, introduced plants were more likely to produce sexual inflorescences and clonal asexual vegetative bulbils, and they invested much more biomass in both reproductive modes. Higher reproductive investment was due to higher proportional allocation of biomass rather than larger plant size. These results are consistent with selection for increased reproduction during range expansion. However, population genetic surveys indicate that recruitment from seed rarely occurs in introduced populations. Hence increased sexual allocation is not an adaptive response to invasion. Although increased clonal reproduction may be advantageous in expanding populations, genetic evidence from introduced populations of B. umbellatus suggests that increased clonal allocation may have arisen via stochastic processes during long-distance transport or a selective filter right at introduction, rather than incremental natural selection during range expansion.     

Reproductive strategy, clonal structure and genetic diversity in populations of the aquatic macrophyte Sparganium emersum in river systems

Many aquatic and riparian plant species are characterized by the ability to reproduce both sexually and asexually. Yet, little is known about how spatial variation in sexual and asexual reproduction affects the genotypic diversity within populations of aquatic and riparian plants. We used six polymorphic microsatellites to examine the genetic diversity within and differentiation among 17 populations (606 individuals) of Sparganium emersum, in two Dutch-German rivers. Our study revealed a striking difference between rivers in the mode of reproduction (sexual vs. asexual) within S. emersum populations. The mode of reproduction was strongly related to locally reigning hydrodynamic conditions. Sexually reproducing populations exhibited a greater number of multilocus genotypes compared to asexual populations. The regional population structure suggested higher levels of gene flow among sexually reproducing populations compared to clonal populations. Gene flow was mainly mediated via hydrochoric dispersal of generative propagules (seeds), impeding genetic differentiation among populations even over river distances up to 50 km. Although evidence for hydrochoric dispersal of vegetative propagules (clonal plant fragments) was found, this mechanism appeared to be relatively less important. Bayesian-based assignment procedures revealed a number of immigrants, originating from outside our study area, suggesting intercatchment plant dispersal, possibly the result of waterfowl-mediated seed dispersal. This study demonstrates how variation in local environmental conditions in river systems, resulting in shifting balances of sexual vs. asexual reproduction within populations, will affect the genotypic diversity within populations. This study furthermore cautions against generalizations about dispersal of riparian plant species in river systems.     

Predominance of clonal reproduction, but recombinant origins of new genotypes in the free-floating aquatic bladderwort Utricularia australis f. tenuicaulis (Lentibulariaceae)

Aquatic plants are a biological group sharing several adaptations to aquatic conditions. The most striking evolutionary convergence in this group is the extensive reliance on clonal reproduction, which largely determines the patterns and process of evolution in aquatic plants. Utricularia australis f. tenuicaulis is a free-floating aquatic bladderwort that reproduces both sexually via seeds and clonally via turions and shoot fragments. Amplified fragment length polymorphism analysis was conducted on 267 ramets collected from 30 populations in Japan. The genotypic diversity within populations was extremely low, regardless of the geographical distribution range: the mean number of genotypes per population (G) was 1.4 and the mean genotypic diversity (D), including monoclonal populations, was 0.17. In contrast to the predominance of a few clones within populations, many of the populations investigated had different genotypes; a large portion of the genetic variation was explained by variation among populations. Character compatibility analysis clearly revealed that somatic mutations did not contribute to the origin of genotypic diversity in this aquatic bladderwort; instead, rare-to-sporadic sexual reproduction probably generated new genotypes. Thus, future studies should examine the role of sexual reproduction in this species from the viewpoint of long-term evolutionary benefits.     

Meiotic recombination dramatically decreased in thelytokous queens of the little fire ant and their sexually produced workers

The little fire ant, Wasmannia auropunctata, displays a peculiar breeding system polymorphism. Classical haplo-diploid sexual reproduction between reproductive individuals occurs in some populations, whereas, in others, queens and males reproduce clonally. Workers are produced sexually and are sterile in both clonal and sexual populations. The evolutionary fate of the clonal lineages depends strongly on the underlying mechanisms allowing reproductive individuals to transmit their genomes to subsequent generations. We used several queen-offspring data sets to estimate the rate of transition from heterozygosity to homozygosity associated with recombination events at 33 microsatellite loci in thelytokous parthenogenetic queen lineages and compared these rates with theoretical expectations under various parthenogenesis mechanisms. We then used sexually produced worker families to define linkage groups for these 33 loci and to compare meiotic recombination rates in sexual and parthenogenetic queens. Our results demonstrate that queens from clonal populations reproduce by automictic parthenogenesis with central fusion. These same parthenogenetic queens produce normally segregating meiotic oocytes for workers, which display much lower rates of recombination (by a factor of 45) than workers produced by sexual queens. These low recombination rates also concern the parthenogenetic production of queen offspring, as indicated by the very low rates of transition from heterozygosity to homozygosity observed (from 0% to 2.8%). We suggest that the combination of automixis with central fusion and a major decrease in recombination rates allows clonal queens to benefit from thelytoky while avoiding the potential inbreeding depression resulting from the loss of heterozygosity during automixis. In sterile workers, the strong decrease of recombination rates may also facilitate the conservation over time of some coadapted allelic interactions within chromosomes that might confer an adaptive advantage in habitats disturbed by human activity, where clonal populations of W. auropunctata are mostly found.     

具混合繁殖策略的草本植物异果舞花姜的居群遗传结构

 居群遗传结构的形成受到各种因素的影响。其中, 繁殖方式可能对居群内遗传变异有极其重要的意义, 而距离隔离也是居群间变异产 生的主要原因之一。异果舞花姜(Globba racemosa)具有混合繁殖策略(以种子进行有性繁殖和以珠芽进行无性克隆繁殖)。调查分布于云南的7 个异果舞花姜居群间有性与无性克隆繁殖的差异。采用ISSR标记研究各个居群的遗传多样性与克隆多样性, 探讨繁殖方式和距离隔离对居群遗 传结构的影响。调查结果表明, 异果舞花姜各个居群存在一定的繁殖差异。ISSR结果显示, 该种在种水平上呈现较高水平的遗传变异 (PPB=71.19%), 大部分的变异来自于居群间(G_ST = 0.590 7)。同时, 异果舞花姜具有较高水平克隆多样性(G/N = 0.88)。遗传多样性和克隆多 样性与繁殖水平的变异间相关性不明显, 说明繁殖方式不是居群遗传结构形成的必要和决定性的因素。居群间的地理距离与遗传距离显著相关 (r = 0.68, p < 0.05), 表明距离隔离是居群间遗传变异形成的重要原因。其它因素(如少量新有性个体的补充、细胞突变、奠基效应等)也对 异果舞花姜居群遗传结构的形成和维持起到了重要作用。     

Consequences of prolonged clonal growth on local and regional genetic structure and fruiting success of the forest perennial Maianthemum bifolium

The balance between clonal propagation and sexual reproduction varies among species. Although theory predicts an impact of clonal growth on both‐ within‐ and between population genetic structure, most empirical evidence available to date does not reveal sharp differences between sexually reproducing and clonal species. This has been attributed mainly to the fact that even low levels of sexual recruitment can maintain high levels of genetic diversity. Here we study the effects of prolonged clonal growth and very low rates of sexual recruitment on the genetic structure of the perennial Maianthemum bifolium, an outcrossing understorey species of temperate forests. Average genotypic diversity (0.37) of the populations, as revealed by AFLP, was above the average values reported for species of similar characteristics, but some populations were extremely poor in genotypes. Fruiting success was positively correlated with genotypic diversity, probably as a result of shortage in mating types and compatible pollen in populations poor in genotypes. This was confirmed by a pollination experiment. Fruiting success increased by a factor three when individuals were hand‐pollinated with pollen from a nearby population compared to hand‐pollinations with pollen from the own population. Furthermore, the fruiting success after natural pollination (control individuals) was positively related to number of nearby populations which could act as pollen sources. Given the limited colonization capacity of the species (no seed flow), and the long time since fragmentation of the forest fragments studied, between‐population genetic differentiation was relatively low (Φ_st=0.14). Lack of genetic drift due to long generation times and very limited sexual recruitment is probably responsible for this. Our results show that prolonged clonal growth and lack of sexual recruitment may affect within‐ and between‐ population genetic structure and the capability for sexual reproduction.     

Clonal growth is enhanced in the absence of a mating morph: a comparative study of fertile stylar polymorphic and sterile monomorphic populations of Nymphoides montana (Menyanthaceae)

Background and Aims

Many aquatic species with stylar polymorphisms have the capacity for clonal and sexual reproduction and are sensitive to the balance of the two reproductive modes when there are a limited number of mating morphs within a population. This study asked how the clonal and sexual reproductive modes perform in populations that contain only a single morph and where fitness gain through sexual reproduction is rare. In clonal aquatic Nymphoides montana, polymorphic populations normally contain two mating morphs in equal frequencies. Populations are sexually fertile and appear to be maintained by pollen transfer between the two partners. However, in a monomorphic population of N. montana where mating opportunities are unavailable, female and male function is impaired and clonality maintains the population. Here, the consequences of intraspecific variation in sexuality were explored between monomorphic and polymorphic N. montana populations in eastern Australia.

Methods

Comparative measurements of male and female fertility, total dry mass and genotypic diversity using ISSR markers were made between populations with variable sexuality.

Key Results and Conclusions

Very few seeds were produced in the monomorphic population under natural and glasshouse conditions due to dysfunctional pollen and ovules. Stigma–anther separation was minimal in the monomorphic population, which may be a consequence of the relaxed selective pressures that regulate the maintenance of sexual function. However, clonal reproduction was favoured at the expense of sexual reproduction in the monomorphic population; this may facilitate the establishment of sterility throughout the population via resource reallocation or pleiotropic effects. The ISSR results showed that the monomorphic population was one large, single genotype, unlike the multi-genotypic fertile polymorphic populations. Evolutionary loss of sex in a clonal population in which a mating morph is absent was evident; under these conditions clonal growth may assure reproduction and expand the population via spreading stolons.     

Ecology of two cytotypes ofButomus umbellatus I. Karyology and breeding behaviour

Chromosome numbers were counted inButomus umbellatus from 99 localities in both Czech and Slovak Republics and the karyotype morphology was studied. Basic chromosome sets are asymmetrical and uniform among diploids (2n=26) and triploids (2n=39). Diploids occur less frequently than triploids in this region. Their clonal populations are usually fertile owing to self-compatibility. The clonal populations of triploids are selfincompatible and usually sterile. Thus, the different self-compatibility is the main biological character distinguishing diploids from triploids. Pollen of triploids is viable in spite of meiotic irregularities in pollen mother cells (PMCs). Hybridization both between cytotypes and between the different triploid genets may take place, if they occur sympatrically. Offspring having diverse chromosome numbers between diploid and triploid level can originate as the consequence of such hybridization.     

Rare sexual reproduction events in the clonal reproduction system of introduced populations of the little fire ant

A unique reproductive system has previously been described in Wasmannia auropunctata, a widespread invasive ant species, where males are produced clonally, female queens are parthenogens, and female workers are produced sexually. However, these findings were mostly based on samples originating from only a limited part of the native range of the species in South America. We used microsatellite markers to uncover the reproductive modes displayed by a large number of nests collected in various invasive W. auropunctata populations introduced 40 years ago into New Caledonia, where the species now forms a single 450-km-long supercolony. Although the main reproduction system in New Caledonia remained clonality for both male and female reproductives, we found evidence of rare sexual reproduction events that led to the production of both new queen and male clonal lineages. All clonal lineages observed in New Caledonia potentially derived from sexual reproduction, recombination, and mutation events from a single female and a single male genotype. Hence, the male and female gene pools are not strictly separated in New Caledonia and the two sexes do not follow independent evolutionary trajectories. Our results also suggest genetic determination for both parthenogenesis and caste. We discuss the evolutionary implications of the emergence of sex in the clonal reproduction system of introduced populations of W. auropunctata.     

The loss of sex in clonal plants

Most plants combine sexual and clonal reproduction, and the balance between the two may vary widely between and within species. There are many anecdotal reports of plants that appear to have abandoned sex for clonal reproduction, yet few studies have quantified the degree of sexual variation in clonal plants and fewer still have determined the underlying ecological and/or genetic factors. Recent empirical work has shown that some clonal plants exhibit very wide variation in sexual reproduction that translates into striking variation in genotypic diversity and differentiation of natural populations. Reduced sexual reproduction may be particularly common at the geographical margins of species' ranges. Although seed production and sexual recruitment may often be limited by biotic and abiotic aspects of the environment in marginal populations, genetic factors, including changes in ploidy and sterility mutations, may also play a significant role in causing reduced sexual fertility. Moreover, environmental suppression of sexual recruitment may facilitate the evolution of genetic sterility because natural selection no longer strongly maintains the many traits involved in sex. In addition to the accumulation of neutral sterility mutations in highly clonal populations, the evolution of genetic infertility may be facilitated if sterility is associated with enhanced vegetative growth, clonal propagation or survival through either resource reallocation or pleiotropy. However, there are almost no experimental data with which to distinguish among these possibilities. Ultimately, wide variation in genotypic diversity and gene flow associated with the loss of sex may constrain local adaptation and the evolution of the geographical range limit in clonal plants.     

FROM CLONAL TO SEXUAL HYBRIDS: GENETIC RECOMBINATION VIA TRIPLOIDS IN ALL-HYBRID POPULATIONS OF WATER FROGS

Speciation via interspecific hybrids is very rare in animals, as compared to plants. Whereas most plants overcome the problem of meiosis between different chromosome sets by tetraploidization, animal hybrids often escape hybrid sterility by clonal reproduction. This comes at the expense of genetic diversity and the ability to purge deleterious mutations. However, here we show that all-hybrid populations of diploid (LR) and triploid (LLR and LRR) water frogs ( Pelophylax esculentus ) have secondarily acquired sexual reproduction. First, in a crossing experiment analyzed with microsatellite markers, triploid hybrids of both sexes and genotypes (LLR and LRR) recombined their homospecific genomes. Second, the great majority of natural populations investigated had low multilocus linkage disequilibrium, indicating a high recombination rate. As predicted from mating system models, the L genome had constant, low levels of linkage disequilibrium, whereas linkage disequilibrium in the R genome showed a significant reduction with increasing proportion of recombining triploids. This direct evidence of sexual reproduction in P. esculentus calls for a change of the conventional view of hybridogens as clonally reproducing diploids. Rather, hybridogens can be independent sexually reproducing units with an evolutionary potential.     

Genotypic diversity revealed by allozymes and oligonucleotide DNA fingerprinting in French populations of the aquatic macrophyte Sparganium erectum

The vast majority of perennial plants reproduce sexually and vegetatively at the same time. This may lead to important variation among clonal plant populations in their amount of genotypic diversity. In order to verify this assumption, we compare the clonal diversity of 10 natural populations of the aquatic clonal macrophyte Sparganium erectum in France. Diversity was quantified by DNA fingerprinting and allozyme electrophoresis for a sample of 10 shoots per population. Two DNA probes (CA)₈ and (TAA)₆ TA, were selected among 10 synthetic oligonucleotide probes containing simple repeat motifs. Both allozymes and DNA fingerprints revealed different amounts of diversity among populations. Five populations consist of a single genotype, whereas two populations were genetically highly diverse. In four of the monomorphic populations, absence of fingerprints diversity was combined with uniformly heterozygous allozyme loci, suggesting that each population was composed of a single clone. In the highly diverse populations, the level of clonal diversity combined with the allele segregation of the two allozyme loci Lap and Est suggests frequent seedling recruitment. The origin of new genotypes remains unclear but the absence of widespread genotypes together with the discrete distribution of Sparganium erectum populations implies that new genotypes are locally produced through sexual reproduction.     

Reduced recombination in gynodioecious populations of a facultative apomictic orchid

Many plants combine sexual reproduction with some form of asexual reproduction to different degrees, and lower genetic diversity is expected with asexuality. Moreover, the ratios of sexual morphs in species with gender dimorphism are expected to vary in proportion to the reproductive success of the sexual process. Hence, sex ratios can directly influence the genetic structure and diversity of a population. We investigated genotypic diversity in 23 populations of a facultative, apomictic gynodioecious orchid, Satyrium ciliatum, to examine the effect on genotypic diversity of variation in the frequency of females and in the amount of sexual reproduction. The study involved one pure female, seven gynodioecious (both females and hermaphrodites present) and 15 hermaphroditic populations. Pollinia receipt was higher in hermaphroditic than in gynodioecious populations. Analyses of variation in ISSRs demonstrated that genotypic diversity was high in all populations and was not significantly different between hermaphroditic and gynodioecious populations. We used character compatibility analysis to determine the extent to which recombination by sexual reproduction contributed to genotypic diversity. The results indicate that the contribution of recombination to genotypic diversity is higher in hermaphroditic than in gynodioecious populations, consistent with the finding that hermaphroditic populations received higher amounts of pollinia. Our finding of reduced recombination in gynodioecious populations suggests that maintenance of sex in hermaphrodites plays an important role in generating genotypic diversity in this apomictic orchid.     

VARIATION IN CONTRIBUTIONS OF ASEXUAL REPRODUCTION TO THE GENETIC STRUCTURE OF POPULATIONS OF THE SEA ANEMONE METRIDIUM SENILE

Northeastern North American populations of the sea anemone Metridium senile show marked differences in levels of genotypic diversity. Comparisons with expectations generated by computer simulation show that some populations are genotypically as diverse as expected for sexually reproducing populations with free recombination, whereas others are significantly less diverse than expected, despite efforts to avoid collecting clonemates. These reductions in diversity are not attributable to the Wahlund effect; they probably result from extensive clonal reproduction. Reduced genotypic diversity may be produced by low rates of recruitment of planktonic larvae, followed by asexual proliferation. The resulting founder effect may account for previously documented random allele-frequency variation between adjacent populations. It is presently uncertain whether the few genotypes found in some populations are particularly well-adapted to local conditions.