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.     

Population Structures in Rhododendron metternichii var. hondoense Assessed with Microsatellites and their Implication for Conservation

Rhododendron metternichii var. hondoense is its great morphological variation. Individuals may have only one erect stem or may have multiple creeping stems, implying that some of them recruit vegetatively. Aims of this study are to ascertain whether a population of R. metternichii var. hondoense consists of clonal plants, and to evaluate relative importance of sexual and asexual recruitments in regard to its conservation. Six microsatellite loci were analyzed in two populations growing in different habitats. One was in a mesic valley consisting of many sprouting and creeping individuals with few seedlings, and the other on a mountain ridge consisting of single stem individuals with many seedlings. Sufficient polymorphisms were found to be present even in the mesic valley population that consisted of many sprouting and creeping stems, indicating that in the past the population was maintained by both sexual and asexual reproduction. The scarcity of seedlings at the mesic valley was due to dense litter cover and low bryophyte mat cover which may be caused by changes in traditional management systems. Required conservation measures are discussed based on these results. Received 8 July 1999/ Accepted in revised form 26 August 1999     

Isolation and characterization of microsatellite loci from Quercus mongolica var. crispula

Microsatellites were isolated and characterized for the Japanese oak species, Quercus mongolica var. crispula, distributed in temperate deciduous forests of Japan. Eleven of the 48 primer pairs designed successfully amplified unambiguous and polymorphic single loci among 67 Q. mongolica var. crispula individuals within a plot in southwestern Japan. The observed and expected heterozygosities of the 11 microsatellite markers ranged from 0.522 to 0.896 and from 0.536 to 0.882, respectively. These polymorphic microsatellite markers are useful for estimating pollen‐mediated gene flow in Q. mongolica var. crispula.     

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.     

Clonal dynamics in western North American aspen (Populus tremuloides)

Clonality is a common phenomenon in plants, allowing genets to persist asexually for much longer periods of time than ramets. The relative frequency of sexual vs. asexual reproduction determines long‐term dominance and persistence of clonal plants at the landscape scale. One of the most familiar and valued clonal plants in North America is aspen (Populus tremuloides). Previous researchers have suggested that aspen in xeric landscapes of the intermountain west represent genets of great chronological age, maintained via clonal expansion in the near absence of sexual reproduction. We synthesized microsatellite data from 1371 ramets in two large sampling grids in Utah. We found a surprisingly large number of distinct genets, some covering large spatial areas, but most represented by only one to a few individual ramets at a sampling scale of 50 m. In general, multi‐ramet genets were spatially cohesive, although some genets appear to be fragmented remnants of much larger clones. We conclude that recent sexual reproduction in these landscapes is a stronger contributor to standing genetic variation at the population level than the accumulation of somatic mutations, and that even some of the spatially large clones may not be as ancient as previously supposed. Further, a striking majority of the largest genets in both study areas had three alleles at one or more loci, suggesting triploidy or aneuploidy. These genets tended to be spatially clustered but not closely related. Together, these findings substantially advance our understanding of clonal dynamics in western North American aspen, and set the stage for a broad range of future studies.     

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.     

Unveiling the origin of Quercus serrata subsp. mongolicoides found in Honshu,Japan, by using genetic and morphological analyses

Quercus mongolica is a tree found in temperate deciduous forests in east Asia. In Japan, Q. mongolica var. crispula is commonly found; moreover, an oak whose morphology is similar to that of Q. mongolica var. mongolica of the Asian continent has been found in certain areas of Honshu and Hokkaido. Recently, the oak found in Honshu was described as Q. serrata subsp. mongolicoides (QSM). However, genetic comparison between this oak and Q. mongolica var. mongolica of the Asian continent has not been performed; the origin of QSM is thus unknown. Therefore, we aimed to determine the origin of QSM by conducting nuclear microsatellite (nSSR), chloroplast DNA (cpDNA) and leaf morphology analyses for the three taxa, as well as other congeners. The cpDNA variation overlapped among the three taxa, suggesting low discrimination ability for these taxa. Although morphological congruency was found between QSM and Q. mongolica var. mongolica, results of nSSR analyses showed that QSM contained a genetic admixture of Q. mongolica var. mongolica of the Asian continent and Q. mongolica var. crispula of Japan, bolstering an admixture hypothesis. The nSSR and cpDNA analyses suggested that Q. mongolica var. crispula can be the progenitor of Q. mongolica var. mongolica and harbors the ancestral cpDNA haplotypes. Therefore, we concluded that QSM might have been created by an admixture that likely occurred within Japan between Q. mongolica var. crispula and putative relict Q. mongolica var. mongolica, which might have diverged in or around Japan from Q. mongolica var. crispula during the late Pleistocene.     

Seedling recruitment in the invasive clonal shrub, Mahonia aquifolium Pursh (Nutt.)

The clonal shrub, Mahonia aquifolium, is an aggressive invader of some forests in central Germany. We analysed the importance of seedling recruitment for the local dynamics of these invasive populations. In contrast to many other clonal plants, repeated seedling recruitment takes place in M. aquifolium, contributing to the colonization of available space within populations. Thus, sexual reproduction is not only important for dispersal to new sites but also for the local invasion␣process. In situations with grass competition (Calamagrostis epigejos) the number of successful recruits is reduced, and clonal growth is the predominant mode of reproduction. Since seedling recruitment results in an increasing number of genets as well as ramets, the investigated populations are still below their carrying capacities. Seedling densities are higher beneath the canopy of adults than outside adult patches, and increase with increasing cover of adult ramets. The spatially clumped pattern of seedling emergence results in density-dependent mortality especially in the most crowded seedling clusters. However, adult neighbourhood imposes a much higher mortality. Therefore, we expect that the importance of seedling recruitment for the dynamics of local M. aquifolium populations will decline as the invasion process continues. Received: 15 January 1996 / Accepted: 28 September 1996     

Causes and consequences of large clonal assemblies in a poplar hybrid zone

Asexual reproduction is a common and fundamental mode of reproduction in plants. Although persistence in adverse conditions underlies most known cases of clonal dominance, proximal genetic drivers remain unclear, in particular for populations dominated by a few large clones. In this study, we studied a clonal population of the riparian tree Populus alba in the Douro river basin (northwestern Iberian Peninsula) where it hybridizes with Populus tremula, a species that grows in highly contrasted ecological conditions. We used 73 nuclear microsatellites to test whether genomic background (species ancestry) is a relevant cause of clonal success, and to assess the evolutionary consequences of clonal dominance by a few genets. Additional genotyping‐by‐sequencing data were produced to estimate the age of the largest clones. We found that a few ancient (over a few thousand years old) and widespread genets dominate the population, both in terms of clone size and number of sexual offspring produced. Interestingly, large clones possessed two genomic regions introgressed from P. tremula, which may have favoured their spread under stressful environmental conditions. At the population level, the spread of large genets was accompanied by an overall ancient (>0.1 Myr) but soft decline of effective population size. Despite this decrease, and the high clonality and dominance of sexual reproduction by large clones, the Douro hybrid zone still displays considerable genetic diversity and low inbreeding. This suggests that even in extreme cases as in the Douro, asexual and sexual dominance of a few large, geographically extended individuals does not threaten population survival.     

Clonal structure of natural populations of Cryptomeria japonica growing at different positions on slopes,detected using RAPD markers

Cryptomeria japonica D. Don (Japanese cedar), an economically important timber species endemic to Japan, is dominant on ridges and upper slopes in cool-temperate natural forests of Kyoto Prefecture. Recruitment of sexual progeny in the area near the Japan Sea is extremely rare, and propagation occurs predominantly through clonal growth by layering. The development pattern that occurs with layering and the resulting complexity of the population structure make it difficult to identify distinct clones, even by excavation. Therefore, we used the randomly amplified polymorphic DNA (RAPD) technique to examine the clonal structure of upper- and lower-slope plots established in two C. japonica populations in Kyoto Prefecture. A total of 263 plants sampled from four plots were analyzed using 10 arbitrarily chosen decamer primers, which produced 50 highly reproducible RAPD bands. There was a different clonal structure in upper- and lower-slope plots. Lower-slope plots were made up of a small number of genets with many ramets, while upper-slope plots were made up of a large number of genets with a few ramets. Clonal diversity measured using PD, Simpson’s D, and Fager’s E was higher in the upper-slope plots. These results show that natural C. japonica populations maintain relatively high clonal variation, compared with other clonal plant species, and that repeated seedling recruitment occurred more frequently in upper-slope plots than in lower-slope plots.     

The effect of herbivores on genotypic diversity in a clonal aquatic plant

In clonal plants, vegetative parts may outcompete seeds in the absence of disturbance, limiting the build‐up of genotypic diversity through repeated seedling recruitment (RSR). Herbivory may provide disturbance and trigger establishment of strong colonizers (seeds) at the expense of strong competitors (clonal propagules). In the clonal aquatic fennel pondweed Potamogeton pectinatus, two distinct herbivore guilds may modify the dynamics of propagation. In winter, Bewick's swans may deplete patches of tubers, promoting seedling establishment in spring. In summer, seed consumption by waterfowl can reduce the density of viable seeds but grazing may also reduce tuber production and hence facilitate seedling establishment. This study is among the first to experimentally test herbivore impact on plant genotypic diversity. We assess the separate and combined effects of both herbivore guilds on genotypic diversity and structure of fennel pondweed beds. Using microsatellites, we genotyped P. pectinatus from an exclosure experiment and assessed the contribution of herbivory, dispersal and sexual reproduction to the population genetic structure. Despite the predominance of clonal propagation in P. pectinatus, we found considerable genotypic diversity. Within the experimental blocks, kinship among genets decreased with geographic distance, clearly identifying a role for RSR in the maintenance of genotypic diversity within the fennel pondweed beds. However, over a period of five years, none of the herbivory treatments affected genotypic diversity. Hence, sexual reproduction on a local scale is important in this putatively clonal plant and possibly sufficient to ensure a relatively high genotypic diversity even in the absence of herbivores. Although we cannot preclude a role of herbivory in shaping genotypic diversity of a clonal plant, after five years of exclusion of the two investigated herbivore guilds no measurable effect on genotypic diversity was detected.     

Gnawing damage by rodents to the seedlings ofFagus crenata andQuercus mongolica var.grosseserrata in a temperateSasa grassland-deciduous forest series in southwestern Japan

The effects of dwarf bamboo,Sasa, cover on the initial morrality of hardwood seedlings were investigated by transplanting 1-year-old beech (Fagus crenata) and current-year oak (Quercus mongolica var.grosseserrata) seedling to three different stands; old-growth beech and secondary oak forests withSasa undergrowth, and aSasa grassland in a grassland-forest series near the top of Mt Jippo, southwestern Japan. The most frequent cause of seedling morrality was gnawing of the stems by rodents. In the beech forest, the gnawing was more likely to occur underSasa cover, suggesting that it provides a good habitat for rodents on the beech forest floor. TheSasa under growth may thus play an imporrant role in regeneration of beech forest. In the oak floor, mortality of both species was low and only a little gnawing occurred during a year. However, no natural oak seedling were found in the forest even after a mast year. This may be because most of the acorns disappeated before establishment. The early-stage demography of hardwood seedling as oak may thus play an imporrant role in regeneration of oak forest. In theSasa grassland where the seed supply is small, almost all of the seedlings died fromo gnawing regardless of the presence ofSasa cover. These factors prevent the recruitment of a sizable seedling bank. Rodents may thus play an imporrant role in maintenance of theSasa grassland.     

Vegetative multiplication and patch colonisation of Asarum europaeum subsp. europaeum L. (Aristolochiaceae) inferred by a combined morphological and molecular study

Asarum europaeum subsp. europaeum (Aristolochiaceae) is a rhizomatous herb forming distinct patches in calcareous broadleaved forests. Within natural stands, patches were mapped. In two regions, at least four patches were dug out, and connections between leaf modules through rhizomatous spacers were checked for signs of clonal reproduction (decay, breaking). Modules were sampled for amplified fragment length polymorphism (AFLP) fingerprinting to test whether they represent unique genets or are merigenets of a larger genet (split up by clonal reproduction), respectively.Morphologically, merigenet-relationships were only revealed in few cases with disrupted spacers between modules. With the obtained AFLP profiles for two primer combinations, the samples could be assigned to genets; clonal descendants of the same genet were readily identified. In one patch analysed in detail, 18 samples from 17 unconnected “plants” belonged to only two genets, which were morphologically divided into two and 15 merigenets, respectively. These two genets probably belonged to different maternal lineages and came into contact after lateral spread from the established clones. They showed divergent affinities to samples from adjacent patches (which all represented unique genets).The findings support the suitability of the combined morpho-ecological and molecular approach: compared to either method alone, it allows a more detailed analysis and interpretation of the fine-scale clonal structure, patch colonisation and especially of vegetative multiplication (with morpho-ecological studies to discern clonal growth and clonal reproduction and AFLP fingerprinting for genet and merigenet identification, respectively).     

CONTRIBUTIONS OF SEXUAL AND ASEXUAL REPRODUCTION TO POPULATION STRUCTURE IN THE CLONAL SOFT CORAL,ALCYONIUM RUDYI

Numerous studies of population structure in sessile clonal marine invertebrates have demonstrated low genotypic diversity and nonequilibrium genotype frequencies within local populations that are monopolized by relatively few, highly replicated genets. All of the species studied to date produce planktonic sexual propagules capable of dispersing long distances; despite local genotypic disequilibria, populations are often panmictic over large geographic areas. The population structure paradigm these species represent may not be typical of the majority of clonal invertebrate groups, however, which are believed to produce highly philopatric sexual propagules. I used allozyme variation to examine the population structure of the temperate soft coral, Alcyonium rudyi, a typical clonal species whose sexually produced larvae and asexually produced ramets both have very low dispersal capabilities. Like other clonal plants and invertebrates, the local population dynamics of A. rudyi are dominated by asexual reproduction, and recruitment of new sexually produced genets occurs infrequently. As expected from its philopatric larval stage, estimates of genetic differentiation among populations of A. rudyi were highly significant at all spatial scales examined (mean θ = 0.300 among 20 populations spanning a 1100-km range), suggesting that genetic exchange seldom occurs among populations separated by as little as a few hundred meters. Mapping of multilocus allozyme genotypes within a dense aggregation of A. rudyi ramets confirmed that dispersal of asexual propagules is also very limited: members of the same genet usually remain within < 50 cm of one another on the same rock surface. Unlike most previously studied clonal invertebrates, populations of A. rudyi do not appear to be dominated by a few widespread genets: estimates of genotypic diversity (G_o) within 20 geographically distinct populations did not differ from expectations for outcrossing, sexual populations. Despite theoretical suggestions that philopatric dispersal combined with typically small effective population sizes should promote inbreeding in clonal species, inbreeding does not appear to contribute significantly to the population structure of A. rudyi. Genet genotype frequencies conformed to Hardy-Weinberg expectations in all populations, and inbreeding coefficients (f) were close to zero. In general, the population structure of A. rudyi did not differ significantly from that observed among outcrossing sexual species with philopatric larval dispersal. Age estimates suggest, however, that genets of A. rudyi live for many decades. Genet longevity may promote high genotypic diversity within A. rudyi populations and may be the most important evolutionary consequence of clonal reproduction in this species and the many others that share its dispersal characteristics.     

Establishment and genetic structure of Empetrum hermaphroditum populations in northern Sweden

Abstract. In late successions of the boreal forest of northern Sweden the evergreen dwarf shrub Empetrum hermaphroditum forms an extensive cover and is believed to spread mainly vegetatively through layering. To analyse the process of population establishment and the relative importance of sexual vs asexual reproduction and the spatial clonal distribution of this species we selected one mainland and two island sites of different post‐fire successional ages (145, 375 and 1720 yr since last fire, respectively). Using 61 polymorphic RAPD markers, we found 96 genotypes in a total of 133 samples. All three populations showed high levels of genetic variation. AMOVA analysis revealed that 33% of the total variation resided among sites, 26% resided among plots within sites and 41% was due to variation within plots. The youngest population had only 14% clonal fraction. In contrast, the oldest population had > 30% clonal fraction and many genets had dimensions of 10–40 m and were intermingled. It appears that E. hermaphroditum establishes by seeds to a larger extent than previously thought and that the clonal spread by layering is rather slow.     

Optimum reproduction and dispersal strategies of a clonal plant in a metapopulation: a simulation study with Hieracium pilosella

Clonal spread is favoured in many plants at the expense of seed production in order to expand rapidly into open habitats or to occupy space by forming dense patches. However, for the dynamics of a population in a patchy landscape seed dispersal remains important even for clonal plants. We used a spatially explicit individual-based metapopulation model to examine the consequences of two trade-offs in Hieracium pilosella L: first, between vegetative and sexual reproduction, and second, between short and far-distance dispersal of seeds. Our main question was, what are the environmental conditions that cause a mixed strategy of vegetative and sexual reproduction to be optimal. The model was parameterised with field data on local population dynamics of H. pilosella. Patch dynamics were given firstly by disturbance events that opened patches in a matrix of a clonal grass that were colonisable for H. pilosella, and secondly by the gradual disappearance of H. pilosella patches due to the expanding grass. Simulations revealed opposing selection pressures on traits determined by the two trade-offs. Vegetative reproduction is favoured by local dynamics, i.e. the need for maintenance and expansion of established populations, whereas seed production is favoured by the necessity to colonise empty habitats. Similar pressures act on the proportion of seeds dispersed over short and far distances. Optimum reproductive and dispersal strategies depended on habitat quality (determined by seedling establishment probability), the fraction of dispersed seeds, and the fraction of seeds lost on unsuitable ground. Under habitat conditions supporting moderate to low seedling establishment, between 20% and 40% of reproductive effort in H. pilosella should be devoted to sexual reproduction with at least 10% of the seeds dispersed over distances suitable to attain empty patches. We conclude that in a spatially heterogeneous landscape sexual seed production in a clonal plant is advantageous even at the expense of local vegetative growth.     

High variation in clonal vs. sexual reproduction in populations of the wild strawberry, Fragaria virginiana (Rosaceae)

Background and Aims

Many plants reproduce both clonally and sexually, and the balance between the two modes of reproduction will vary among populations. Clonal reproduction was characterized in three populations of the wild strawberry, Fragaria virginiana, to determine the extent that reproductive mode varied locally between sites. The study sites were fragmented woodlands in Cook County, Illinois, USA.

Methods

A total of 95 strawberry ramets were sampled from the three sites via transects. Ramets were mapped and genotyped at five variable microsatellite loci. The variability at these five loci was sufficient to assign plants to clones with high confidence, and the spatial pattern of genets was mapped at each site.

Key Results

A total of 27 distinct multilocus genotypes were identified. Of these, 18 genotypes were detected only once, with the remaining nine detected in multiple ramets. The largest clone was identified in 16 ramets. No genets were shared between sites, and each site exhibited markedly different clonal and sexual recruitment patterns, ranging from two non-overlapping and widespread genets to 19 distinct genets. Only one flowering genet was female; the remainder were hermaphrodites.

Conclusions

Local population history or fine-scale ecological differences can result in dramatically different reproductive patterns at small spatial scales. This finding may be fairly widespread among clonal plant species, and studies that aim to characterize reproductive modes in species capable of asexual reproduction need to evaluate reproductive modes in multiple populations and sites.Key words: Clonal structure, gynodioecy, Fragaria virginiana, microsatellites, population genetic structure     

Contrasting patterns of clonality and fine-scale genetic structure in two rare sedges with differing geographic distributions

For plants with mixed reproductive capabilities, asexual reproduction is more frequent in rare species and is considered a strategy for persistence when sexual recruitment is limited. We investigate whether asexual reproduction contributes to the persistence of two co-occurring, rare sedges that both experience irregular seed set and if their differing geographic distributions have a role in the relative contribution of clonality. Genotypic richness was high (R=0.889±0.02) across the clustered populations of Lepidosperma sp. Mt Caudan and, where detected, clonal patches were small, both in ramet numbers (⩽3 ramets/genet) and physical size (1.3±0.1 m). In contrast, genotypic richness was lower in the isolated L. sp. Parker Range populations, albeit more variable (R=0.437±0.13), with genets as large as 17 ramets and up to 5.8 m in size. Aggregated clonal growth generated significant fine-scale genetic structure in both species but to a greater spatial extent and with additional genet-level structure in L. sp. Parker Range that is likely due to restricted seed dispersal. Despite both species being rare, asexual reproduction clearly has a more important role in the persistence of L. sp. Parker Range than L. sp. Mt Caudan. This is consistent with our prediction that limitations to sexual reproduction, via geographic isolation to effective gene exchange, can lead to greater contributions of asexual reproduction. These results demonstrate the role of population isolation in affecting the balance of alternate reproductive modes and the contextual nature of asexual reproduction in rare species.     

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.     

Interspecific variation in clonality in temperate lianas revealed by genetic analysis: Do clonal proliferation processes differ among lianas?

Lianas are important components of forest communities with a considerable impact on overall forest structure and function. Lianas are characterized by extensive clonal reproduction on the forest floor, which is important for their establishment and growth. Previous studies have suggested that clonal reproduction strategies vary substantially among liana species; however, few studies have quantitatively evaluated the clonality strategy in multiple co-occurring liana species. The primary objective of the present study was to evaluate the relative contribution of clonal reproduction and to understand the clonal proliferation processes in co-occurring liana species by assessing both small stems on the forest floor and mature stems that climbed trees. The clonal reproduction strategy in four common liana species (stem twiner: Wisteria floribunda; root climbers: Schizophragma hydrangeoides, Euonymus fortunei and Rhus ambigua) in a 6-ha plot and a belt transect within an old-growth temperate forest in central Japan was evaluated using genetic analysis. The contribution of clonal reproduction was smaller in root climbers than in W. floribunda. All W. floribunda genets with small ramets in the understory had genetically identical ramets that climbed trees, whereas few such ramets were found in root climbers. This indicates that W. floribunda mature ramets laterally produce small ramets via clonal proliferation, whereas seedlings of root climbers grow horizontally to climb trees. The results indicate that the clonal reproduction processes differ greatly among lianas and the clonal growth in lianas plays a contrasting role in their life-history strategy.