Clonal reproduction, vegetative multiplication and habitat colonisation in Tussilago farfara (Asteraceae): A combined morpho-ecological and molecular study

Using a combined morpho-ecological and molecular (amplified fragment length polymorphism (AFLP) fingerprinting) approach, clonal growth, clonal reproduction and the resulting vegetative multiplication were examined in the pioneer species Tussilago farfara (Asteraceae).

In two forest plots in Berlin (Germany), all modules and spacers of the plant were mapped and dug out. The plants showed intense clonal growth with leaf modules connected through long branched rhizomes. The rhizomes were brittle and regularly exhibited rotting zones and breaks. This clonal reproduction can occur rather fast; in a submersed stand in Lanke (Brandenburg, Germany), rhizomes showed signs of disintegration only a few weeks after the modules developed. As a consequence, modules become separated and genets fragment into merigenets.

Using AFLP fingerprints, the relationships of samples from the two dug out plots were assessed: 15 and 18 analysed samples (representing 13 and 16 merigenets) belonged to only two and three distinct genets, respectively. Each of these genets was split into at least two to eleven morphologically independent merigenets, indicating frequent and effective clonal reproduction.

The study highlights the relevance of vegetative multiplication in T. farfara, achieved rapidly through repeated clonal growth and subsequent clonal reproduction. The typical strategy of habitat colonisation and maintenance is described and illustrated for this pioneer species.     

Clonal growth and its effects on male and female reproductive success in <Emphasis Type="Italic">Prunus ssiori</Emphasis> (Rosaceae)

Clonal growth can increase not only floral display but also geitonogamy and may affect sexual reproduction both positively and negatively. A clonal woody species, Prunus ssiori, was partially self-incompatible according to a pollination experiment. Its main pollinators, bumble bees, were often observed to consecutively visit inflorescences within a tree. Clone identification revealed that its genets formed mutually exclusive patches. These features suggest frequent geitonogamous pollination. In a 6.24-ha plot, 212 trees belonged to 59 genets, and 42 genets consisted of a single tree, whereas the rest contained two or more clonal trees. The largest genet had 65 trees and occupied 0.4 of a hectare. Fruit set was measured in 127 inflorescences sampled from nine maternal trees at the center of the plot. Paternal genets of 107 of their 300 seeds were assigned in the plot using microsatellites. There were no selfed seeds. Male reproductive success (the probability that individual trees of each genet sired a seed) increased as tree size increased, as the distance between the trees and maternal trees decreased, when the genet did not contain the maternal trees, and when the genet consisted of a single tree. Female reproductive success (fruit set in individual inflorescences of each maternal tree) increased as the within-tree geitonogamy index, which reflected the frequency of pollination within the maternal tree, decreased. These results suggest that clonal growth reduces male reproductive success, at least, in P. ssiori, because of pollen discounting.     

Fine-scale clonal structure and diversity within patches of a clone-forming dioecious shrub, Ilex leucoclada (Aquifoliaceae)

BACKGROUND AND AIMS: The mode of reproduction (sexual vs. asexual) is likely to have important effects on genetic variation and its spatial distribution within plant populations. An investigation was undertaken of fine-scale clonal structure and diversity within patches of Ilex leucoclada (a clone-forming dioecious shrub). METHODS: Six patches were selected in a 1-ha plot previously established in an old-growth beech forest. Two of the selected patches were composed predominantly of stems with male flowers (male patch), and two contained stems with predominantly female flowers (female patch). The remaining two patches contained stems with male flowers and stems with female flowers in more or less equal proportions (mixed patch). Different genets were distinguished using random amplified polymorphic DNA (RAPD) markers. KEY RESULTS: One hundred and fifty-six genets with different RAPD phenotypes were identified among 1928 stems from the six patches. Among the six patches, the male patches had the lowest clonal diversity, and the mixed patches had the highest. Distribution maps of the genets showed that they extended downhill, reflecting natural layering that occurred when stems were pressed to the ground by heavy snow. In every patch, there were a few large genets with many stems and many small genets with a few stems. CONCLUSION: The differences in clonal diversity among patches may be due to differences in seedling recruitment frequencies. The skewed distribution of genet size (defined as the number of stems per genet) within patches may be due to differences in the timing of germination, or age (with early-establishing genets having clear advantages for acquiring resources) and/or intraspecific competition.     

Reproductive demography of ramets and genets in a rhizomatous clonal plant <Emphasis Type="Italic">Convallaria keiskei</Emphasis>

Clonal growth occurring below the ground makes it difficult to identify individuals and demonstrate the demographic features of a focal plant species. In this study, genotypically identified ramets of a rhizomatous clonal herb, Convallaria keiskei Miq., were monitored for their growth, survival, and reproduction from 2003 to 2006. After the monitoring period, their subterranean organs were excavated to explore the underground connections of established ramets and the direction of clonal growth. We then combined data on the fate of the monitored ramets with the information of rhizome connections, clarifying reproductive demography at both the ramet and genet levels. Although each ramet initiated both sexual reproduction (via flowering) and clonal growth, clonal growth tended to precede sexual reproduction. In a surveyed genet, 51.0% of ramets produced flowers and 29.6% generated clonal offspring during the study period. Consequently, we clarified the reproductive demography of C. keiskei: clonal growth tended to precede flowering in a ramet, and a genet can keep reproducing every season at the genet level, despite a ramet not having inflorescence every year.     

Clonal spread of invasive Ludwigia hexapetala and L. grandiflora in freshwater wetlands of California

Ludwigia hexapetala and L. grandiflora are recent, aggressive invaders of freshwater wetlands in California. To assess the relative role of sexual versus clonal reproduction in invasive spread, we used AFLP markers to genotype 794 ramets of L. hexapetala sampled from 27 populations in three watersheds of California, and 150 ramets of L. grandiflora from five populations in a fourth watershed. We then used two analytical methods, similarity thresholds and character incompatibilities, to distinguish genotypic variation within genets (clones) from variation between genets. Our analyses revealed extremely limited genotypic and genet variation in invasive L. hexapetala and L. grandiflora within California. Within L. hexapetala, 95% of the ramets analyzed represented a single genet. The genet was the only one detected in 20 populations. The remaining seven populations contained two to nine genets. Within L. grandiflora, all ramets were of only one genotype. Thus, invasive spread within and between populations, and across watersheds, appears to be almost exclusively clonal and brought about by the dispersal of vegetative propagules. The extremely low seedling recruitment indicates that management should target vegetative dispersal and growth.     

Vegetative reproduction and clonal diversity in Rhytidium rugosum (Rhytidiaceae, Bryopsida) inferred by morpho-anatomical and molecular analyses

Many bryophytes exhibit specific life forms that result in the formation of distinct patches. This is primarily achieved by consequent vegetative multiplication through indeterminate growth and vegetative reproduction. Little, however, is known about the underlying mechanisms and genetic patterns. In this study on vegetative multiplication in bryophytes, we apply morpho-anatomical and molecular (AFLP fingerprinting) techniques to analyze the vegetative reproduction system, clonal diversity, and habitat colonization of a pleurocarpous moss, Rhytidium rugosum (Rhytidiaceae). Morpho-anatomically, three types of vegetative diaspores are identified and illustrated: ramets, separated after decay and disintegration of older shoot parts (clonal reproduction); brood branches/branchlets; and caducous shoot apices (both vegetative reproduction s.str.). The AFLP fingerprinting of 35 samples (from two plots in Thuringia, further German populations, and from France, Russia, and Canada) resulted in the identification of three clones from the two plots, each comprising two to 15 samples with identical fingerprints. Samples from one clone occurred in both plots, thus proving dispersal of vegetative diaspores. The AFLP analysis further revealed close relationships of the plot samples, which suggest a clonal rather than generative (sexual) origin of the population.     

Intraspecific competition and light effect on reproduction of Ligularia virgaurea,an invasive native alpine grassland clonal herb

The relationship between sexual reproduction and clonal growth in clonal plants often shows up at the ramet level. However, only a few studies focus on the relationship at the genet level, which could finally account for evolution. The sexual reproduction and clonal growth of Ligularia virgaurea, a perennial herb widely distributed in the alpine grasslands of the Qinghai‐Tibetan Plateau of China, were studied under different competition intensities and light conditions at the genet level through a potted experiment. The results showed that: (1) sexual reproduction did not depend on density or light, and increasing clonal growth with decreasing density and increasing light intensity indicated that intraspecific competition and light intensity may affect the clonal life history of L. virgaurea; (2) both sexual reproduction and clonal growth show a positive linear relationship with genet size under different densities and light conditions; (3) a threshold size is required for sexual reproduction and no evidence of a threshold size for clonal growth under different densities and light conditions; (4) light level affected the allocation of total biomass to clonal and sexual structures, with less allocation to clonal structures and more allocation to sexual structures in full sunlight than in shade; (5) light determined the onset of sexual reproduction, and the genets in the shade required a smaller threshold size for sexual reproduction to occur than the plants in full sunlight; and (6) no evidence was found of trade‐offs between clonal growth and sexual reproduction under different densities and light conditions at the genet level, and the positive correlation between two reproductive modes indicated that these are two integrated processes. Clonal growth in this species may be viewed as a growth strategy that tends to maximize genet fitness.     

Does sex make a difference? Genetic diversity and spatial genetic structure in two co-occurring species of Gagea (Liliaceae) with contrasting reproductive strategies

Gagea lutea and G. spathacea are spring geophytes naturally co-occurring in woodlands, characterised by contrasting reproductive strategies probably caused by divergent ploidy levels. The hexaploid G. lutea relies on vegetative reproduction by subterranean bulbils in young stages but completely switches to sexual reproduction once a certain bulb size is attained. The nonaploid G. spathacea seems to be sterile and reproduces only vegetatively; the plants continue to form bulbils even in the rare event of flowering. This study used AFLP genotyping to investigate the consequences of these reproductive strategies for genetic diversity. For 150 and 100 samples from three Western Pomeranian populations of G. lutea and G. spathacea, respectively, AFLP fingerprints were analysed for three different spatial scales, the patch, the transect, and the region. Applying a threshold for genotypic identity of <0.05 simple matching distance, 22?C30 genets were detected in the three G. lutea populations, with all genets confined to single populations. Clonal genets consisted of 2?C9 samples and extended over up to 28?m, but never occupied the whole length of a transect; 67?C75% of all patches had different genets. Genetic distances between genets within populations were similar to those recorded between populations. Genotyping of G. spathacea revealed a single clonal genet for all three populations sampled within a distance of 30?km. The absent genetic diversity confirms the suspected sexual sterility. Gagea spathacea seems to be one of the few non-apomictic, fully clonal vascular plants able to occupy a significant range solely by dispersal of vegetative diaspores.     

Clonal Diversity in Differently-Aged Populations of the Pseudo-Annual Clonal Plant Circaea lutetiana L.

Abstract: In many clonal plant species seedling recruitment is restricted to short colonization episodes early in the development of the population, and clonal diversity (i.e., genet diversity) in the population is expected to decrease with increasing population age. In established populations of the pseudo-annual Circaea lutetiana seedling recruitment has previously not been observed. Therefore, we expected established populations to have low clonal diversities. We analysed number and frequency of genets and spatial distribution of genets in six differently-aged C. lutetiana populations with the use of four informative RAPD primers. We found relatively low clonal diversities in young populations but very high clonal diversities in established populations. Therefore, the hypothesis was rejected that seedling recruitment does not occur in established populations. Moreover, we did not find large genet size asymmetries in established populations. Genet size differences can be caused by stochastic processes or by fitness related traits, such as differences in vegetative reproduction. Because vegetative propagation of ramets is dependent on ramet size, and the number of ramets and the size of each ramet determine genet size, we expected that large genets produced, on average, large ramets. However, this was not the case, suggesting that stochastic processes caused genet size differences. Genet size may also be bounded if spatial distribution of genets is affected by micro-habitat differences. For this we expected to find a clumped spatial distribution of ramets of the same genet. However, ramets of large genets were always found intermingled with ramets belonging to other genets.     

Infertile times: response to damage in genets of the clonal sedge Carex bigelowii

Tolerance to grazing is a plant trait that can be adaptive in systems where plants are subjected to a diversity of herbivore attack types. To test the tolerance ability of the clonal sedge Carex bigelowii, which is food plant to several herbivores in alpine and arctic areas, and the potential fitness costs of this tolerance, replicated units of genets were subjected to three levels of damage throughout three consecutive seasons. The three levels of treatment were no damage, light damage and heavy damage, and the damage was conducted by tearing off all plant material at 3 and 0 cm above-ground respectively. The genets had no tolerance under damage in terms of sexual reproduction. In terms of clonal reproduction the genets showed tolerance under light damage but not under heavy damage. However, no fitness cost was found for this tolerance ability, i.e. genets had higher reproduction and growth under no damage. The average ramet weight had a similar decrease under both a low and high damage treatment. Changed partitioning of biomass between plant parts and reduced concentration of total non-structural carbohydrates (TNC) in storage organs are possible mechanisms for the ability to uphold clonal reproduction in response to damage. There were no significant indications that tolerance ability or its fitness cost differed between genets. Our results suggest that when subjected to heavy damage genets will only reproduce vegetatively. Consequently, it seems C. bigelowii has evolved to allocate resources to the survival of an already successful genet rather than to a potential new genet of unknown success.     

Clonal structure and genet-level sex ratios suggest different roles of vegetative and sexual reproduction in the clonal moss Hylocomium splendens

The allozyme haplotype was determined for 157 ramets of the unisexual, perennial, clonal moss Hylocomium splendens within five 10×10 cm plots, which had been the subject of demographic studies over a 5-yr period. In addition, 25 shoots were analyzed from outside the plots and from four neighbouring patches. Only four haplotypes were encountered within the plots; one female type occurred in all plots and one male type in four plots, whereas two male haplotypes occurred in only one plot. Genets grew intermingled in all but one plot. The sex ratio within the five plots was female-biased at the ramet level (male:female=1:2.6), but male-biased at the genet level (3:1). Sporophytes were produced abundantly during the study period, but no signs of recruitment from spores were observed in the plots. Nine additional genets were encountered in neighbouring patches but from only one patch each. Four (44%) of these could potentially have been derived from spores generated within the plots. Our results suggest that each patch of H. splendens is colonized by a small number of genets, whereas different patches have different sets of genets, i.e. clonal diversity is determined by vegetative reproduction at within-patch scales and structured by sexual processes at among-patch scales.     

Dynamics of distribution and performance of ramets constructing genets: a demographic�Cgenetic study in a clonal plant, Convallaria keiskei

Background and Aims

In clonal plants producing vegetative offspring, performance at the genet level as well as at the ramet level should be investigated in order to understand the entire picture of the population dynamics and the life history characteristics. In this study, demography, including reproduction and survival, the growth patterns and the spatial distributions of ramets within genets of the clonal herb Convallaria keiskei were explored.

Methods

Vegetative growth, flowering and survival of shoots whose genets were identified using microsatellite markers were monitored in four study plots for 3 years (2003–2005). The size structures of ramets in genets and their temporal shifts were then analysed. Their spatial distributions were also examined.

Key Results

During the census, 274 and 149 ramets were mapped in two 1 × 2 m plots, and 83 and 94 ramets in two 2 × 2 m quadrats. Thirty-eight genotypes were identified from 580 samples. Each plot included 5–18 genets, and most ramets belonged to the predominant genet(s) in each plot. Shoots foliated yearly for several years, but flowering ramets did not have an inflorescence the next year. A considerable number of new clonal offspring persistently appeared, forming a bell-shaped curve of the size structure of ramets in each genet. Comparing the structures modelled by the normal distributions suggested variation among ramets belonging to a single genet and variation among genets. Furthermore, spatial analyses revealed clumped and distant distributions of ramet pairs in a genet, in which the distant patterns corresponded to the linearly elongating clonal growth pattern of this species.

Conclusion

Characteristics of ramet performances such as flowering and recruitment of clonal offspring, in addition to growth, played a large part in the regulation of genet dynamics and distribution, which were different among the studied genets. These might be characteristics particularly relevant to clonal life histories.Key words: Clonal plant, Convallaria keiskei, demography, genet, genetic identification, growth pattern, life history, ramet, spatial distribution     

Clonal organization of populations of Asarum europaeum and Maianthemum bifolium in contrasting woodland habitats

Populations of two rhizomatous species, Asarum europaeum (asarabacca) and Maianthemum bifolium (May lily), were examined in two, and four forest habitats respectively, in the Roztocze National Park (south-eastern Poland). May lily populations were studied in habitats: the Carpathian beechwood, upland mixed fir forest, subboreal moist mixed coniferous forest and bog-alder forest. Asarabacca was studied in two habitats: beechwood and Scots pine community (an 80-year-old plantation). In both the species studied intra- and inter-populational differences of the size of genets in terms of above- and below-ground parts of individuals as well as the biomass and area occupied were observed. In May lily populations the greatest mean number of shoots per genet was found in the fir forest (11.62±3.29), a value almost twice as great as that in the moist coniferous forest and nearly three times greater than in the bog-alder forest. Total rhizome length was also the greatest in the fir forest (351.9±98.7 cm) followed by moist coniferous forest, beechwood and alder forest habitats. In all populations of May lily a greater part of total dry weight biomass is in below-ground organs. The greatest biomass value of a genet was found in the fir forest (4.275 g), the smallest in the bog-alder forest (0.110 g). All populations differed significantly in terms of leaf area, leaf length (with the exception of fir forest and beechwood habitats where the values were the greatest), and leaf width (excluding moist coniferous and bog-alder forests which had the smallest values). In the case of asarabacca, both the mean number of ramets per genet (3.36±0.45 vs. 2.49±0.20) and total rhizome length (40.3±6.4 cm vs. 21.1±1.8 cm) were greater in the beechwood habitat than in the pine community. In the first population genets had 3–5 times greater the total biomass of those from the pine community. Only genets of the latter had proportionately more dry weight biomass in above-ground parts. It seems to be correlated with greater rhizome dieback and disintegration of genets into smaller units. Both populations were significantly different in terms of all examined parameters of leaves. Genets of both the species studied were found to have their own structure of developmental phases that often differed for shoots and rhizomes.     

Patch establishment and development of a clonal plant,Polygonum cuspidatum,on Mount Fuji

Microsatellite analysis was used to investigate the patch establishment and development of Polygonum cuspidatum Sieb. et Zucc, a clonal herbaceous plant that dominates the primary succession on the southeast slope of Mount Fuji. Genotypes of P. cuspidatum in 155 patches at the study site differed from each other. This indicates that P. cuspidatum patches are initially established by seed dispersed on the bare scoria field, and not by clonal rhizome extension. Genetic differentiation was estimated using the FST values between subpopulations at the study site. There was almost no genetic differentiation between subpopulations, indicating the presence of massive gene flow. The pollen fathers of seeds and maternal genets of current-year seedlings were inferred from the microsatellite allele composition by a simple exclusion method. The wide, random distribution of pollen fathers suggests that pollen dispersal occurs over a broad area. Maternal analysis showed a tendency for seed dispersal to be biased to the area nearby and down slope from the mother plants. Patch establishment under massive gene flow may result from such pollen and seed dispersal. To understand the process of patch development, aerial photographs taken from 1962 to 1999 were compared, and then genets in each of 36 patches were identified from the microsatellite genotypes of P. cuspidatum shoots. The comparison of aerial photographs showed that most of the patches enlarged each year and that some neighbouring patches combined during growth. Genet analysis demonstrated a high correlation between patch area and the area of the largest genet within it, and that new genets were recruited at the patch periphery. These findings indicate that both vegetative and sexual reproduction, i.e. rhizome extension and the establishment of new seedlings, contribute to the development of P. cuspidatum patches.     

Genet structure and determinants of clonal structure in a temperate deciduous woodland herb, Uvularia perfoliata

1 We used isozyme variation to examine the genet structure of Uvularia perfoliata patches in gap and closed canopy habitats in a temperate deciduous forest in Maryland, USA.
2 A large patch in a gap habitat was composed of a small number of widely spread genets with many ramets, and a large number of genets with more restricted distribution and few ramets. Genets with many ramets were patchily distributed at a metre scale. Analysis of genet structure on a scale of square centimetres, however, revealed that the genets were highly intermingled with no clear boundaries between them. The presence at both scales of sampling of many genets with unique multilocus genotypes indicated continuing genet recruitment within the population.
3 In the closed canopy habitat, the patches examined were each composed of a single unique multilocus genotype, suggesting that each had developed by asexual propagation following the establishment of a single genet.
4 The clonal structure of U. perfoliata patches in both gap and closed canopy habitats therefore appears to depend on recruitment patterns of genets. Populations in closed canopy habitats are characterized by a 'waiting' strategy, in which asexual ramet production maintains populations until genet recruitment by seed production can occur under the more optimal conditions associated with canopy gaps. Extended sampling suggests that the genetic diversity of U. perfoliata populations is primarily controlled by the disturbance regime of the forest canopy.     

Fitness‐consequences of geitonogamous selfing in a clonal marine angiosperm (Zostera marina)

Plant mating systems have received considerable attention because the proportion of selfed vs. outcrossed progeny is an important evolutionary factor. In clonally reproducing plants, geitonogamous selfing between distant ramets belonging to the same genet is expected to be widespread, yet empirical data are sparse. Nothing is known about between‐ramet selfing in aquatic flowering plants with subaqueous pollen transfer, most of which display pronounced clonal reproduction. From two locations in the western Baltic Sea, I present data on the effects of patch isolation and clonal diversity on the outcrossing rate of eelgrass, Zostera marina L., based on the genotypes of maternal plants and recently fertilized ovules scored at eight microsatellite loci. There were no differences in outcrossing rates between vegetation patches and continuous meadow although patches were nearly always composed of single genets. Quantitative effects of clonal diversity were present in the continuous vegetation where a significant positive correlation between genet diversity and the proportion of outcrossed offspring was detected (Kendall’s τ=0.82, P=0.0017). On a population‐scale as well, the genotypic diversity was positively correlated with outcrossing. The relative fitness of selfed offspring was low (ω ± 95% confidence interval=0.56 ± 0.032 and 0.322 ± 0.15) indicating that geitonogamy incurred substantial fitness costs. Selfing rates in Z. marina may not be in evolutionary equilibrium because of spatial and temporal heterogeneity of clonal size and diversity. The high prevalence of dioecy in seagrasses may have evolved to avoid the fitness costs associated with geitonogamy.     

Genetic structure and reproduction dynamics of Salix reinii during primary succession on Mount Fuji,as revealed by nuclear and chloroplast microsatellite analysis

The early stage of volcanic desert succession is underway on the southeastern slope of Mount Fuji. We used markers of nuclear microsatellites (simple sequence repeats; SSR) and chloroplast microsatellites (cpSSR) to investigate the population genetic structure and reproduction dynamics of Salix reinii, one of the dominant pioneer shrubs in this area. The number of S. reinii genets in a patch and the area of the largest genet within the patch increased with patch area, suggesting that both clonal growth and seedling recruitment are involved in the reproduction dynamics of S. reinii. Five polymorphic cpSSR markers were developed for S. reinii by sequencing the noncoding regions between universal sequences in the chloroplast genome. Nineteen different cpSSR haplotypes were identified, indicating that S. reinii pioneer genets were created by the long-distance dispersal of seeds originating from different mother genets around the study site, where all vegetation was destroyed during the last eruption. Furthermore, the clustered distributions of different haplotypes within each patch or plot suggested that newly colonized genets tended to be generated from seeds dispersed near the initially established mother genets. These results revealed that the establishment of the S. reinii population on the southeastern slope of Mount Fuji involved two sequential modes of seed dispersal: long-distance dispersal followed by short-distance dispersal.     

Female and male fitness consequences of clonal growth in a dwarf bamboo population with a high degree of clonal intermingling

Background and Aims

Although many studies have reported that clonal growth interferes with sexual reproduction as a result of geitonogamous self-pollination and inbreeding depression, the mating costs of clonal growth are expected to be reduced when genets are spatially intermingled with others. This study examined how clonal growth affects both female and male reproductive success by studying a population of a mass-flowering plant, Sasa veitchii var. hirsuta, with a high degree of clonal intermingling.

Methods

In a 10 × 10 m plot, genets were discriminated based on the multilocus genotypes of 11 nuclear microsatellite loci. The relationships between genet size and the components of reproductive success were then investigated. Male siring success and female and male selfing rates were assessed using paternity analysis.

Key Results

A total of 111 genets were spatially well intermingled with others. In contrast to previous studies with species forming distinct monoclonal patches, seed production linearly increased with genet size. While male siring success was a decelerating function of genet size, selfing rates were relatively low and not related to genet size.

Conclusions

The results, in conjunction with previous studies, emphasize the role of the spatial arrangement of genets on both the quantity and quality of offpsring, and suggest that an intermingled distribution of genets can reduce the mating costs of clonal growth and enhance overall fitness, particularly female fitness.     

Plasticity in fitness and fitness-related traits at ramet and genet levels in a tillering grass Panicum miliaceum under patchy soil nutrients

Seedling and root sprout recruitment after a volcanic deforestation was examined in a Prunus ssiori population. According to genotypes of five microsatellite (SSR) loci, 44 genets were identified among 188 ramets ≥ 5 cm DBH in a 2.3-ha plot in 2000. The genets formed mutually exclusive patches. Because only one repeat change in an SSR locus distinguished five genets from adjacent genets, there is a possibility that the five genets derived from somatic mutations as well as sexual reproduction. Thus, at least 39 genets originated from seedlings, and at least 144 ramets derived from root sprouts. The seedlings were recruited after the volcanic eruptions in 1739 and before 1972 because trees of 5 cm DBH were 28 years of age, estimated from annual rings. As the largest ramet recruited from a root sprout was estimated to be 94 years old, root sprout recruitment began between 1739 and 1906. Thus, the estimated minimum rates of recruitment from seedlings and root sprouts were 0.167 y^–1 and 0.618-2.182 y^–1, respectively. The clonal diversity of the P. ssiori population (Simpson's D = 0.92) was higher than that of other clonal plant populations. The inbreeding coefficient was significantly negative (Wright's F _IS = −0.069). These results suggest that the frequent seedling recruitment in an outbreeding system maintains the clonal diversity in the population. This revised version was published online in June 2006 with corrections to the Cover Date.     

Fine-scale spatial structure of genets and sexes in the dioecious plant <Emphasis Type="Italic">Dioscorea japonica</Emphasis>, which disperses by both bulbils and seeds

To understand the evolution of clonal reproduction and the diversity of clonal plants, it is necessary to clarify the characteristics of each clonal habit. There has been little research on whether bulbils alter spatial genetic structure (SGS) because of the lack of connection to maternal ramets. We used simple-sequence-repeat (SSR) markers to determine the fine-scale SGS of the dioecious plant Dioscorea japonica, which disperses both as bulbils and as seeds. We also evaluated the contributions of sexual and clonal reproduction and tested for spatial sex segregation (SSS). We discovered 111 genets from 394 ramets in a 2.8-ha plot. Genotypic richness (R = 0.28) and clonal diversity (Simpson’s D = 0.94, Fager’s E = 0.90) were high. We did not find SSS, suggesting that the population does not suffer from a shortage of mating pairs due to clonal reproduction. The Sp values revealed moderate SGS at the genet level (Sp = 0.013–0.014), and the genets intermingled at a local scale. Significant SGS at the ramet level showed that ramets within the same genet tended to aggregate. We also found a skewed clonal spatial distribution. The spatial extent of genets was positively correlated with the number of ramets within a genet. The contribution of bulbil production to the variance of parent–offspring gene dispersal was about one–fifth the contribution from sexual reproduction. These results suggest that the dispersal via bulbils affects the SGS in D. japonica, although its contribution to gene dispersal is small compared to the contribution of sexual reproduction.