Sexual dimorphism in clonal growth forms and ramet distribution patterns in <Emphasis Type="Italic">Rumex acetosella</Emphasis> (Polygonaceae)

We investigated clonal traits in the dioecious herb Rumex acetosella to characterize sexual dimorphism in clonal forms and to correlate below-ground clonal patterns and above-ground ramet distributions. We recorded creeping root length, branching patterns, ramet and clump (caespitose ramets from the same position on the root) sprouting patterns, and biomass allocations in three females and males. We also estimated the patch size of flowering ramets within a quadrat. No sexual dimorphism was detected in the frequencies of branches and flowering ramets per root length. Male plants allocated proportionally more biomass to below-ground organs. Total root length did not differ between the sexes. Females sprouted more clumps with fewer flowering ramets per root length than males, which sprouted fewer clumps with more flowering ramets, which meant that clump sprouting patterns were phalanx-like in females and guerrilla-like in males. Flowering ramets were aggregately distributed in both females and males and patch sizes were similar between sexes, indicating that the spreader propagations were not found in the guerrilla-like males. We assumed that sexual dimorphism occurred in response to physiological integration for higher reproductive effort in females.     

Impact of Culm Harvest on Seed Production in a Monocarpic Bamboo

Successful regeneration of bamboos from seed is a key issue in the ecology of many tropical regions and the livelihoods of their inhabitants. The gregarious monocarpy of many bamboos may be driven by a need to satiate seed predators by seeding in abundance at infrequent intervals. In long‐lived clonal monocarps, seed production is expected to be positively related to the success of the clone in generating more and larger ramets during its lifetime. Ramification may be constrained by harvesting of culms, but it is unclear whether the reduction in productivity is proportional to the loss of reproductive biomass. We counted the seed produced by 661 culms (ramet stems) sampled from 90 clumps of the gregariously monocarpic bamboo Schizostachyum dullooa that is intensively harvested by villagers in northeastern India. The smallest clumps had fewer culms and few or no culms more than one year old. Seed production was indeed positively related to culm size and the number of culms in a clump. First‐year culms were markedly more productive than older culms after controlling for culm diameter and clump size. There was a negative effect of clump size on productivity per culm which may occur because clumps that had been harvested heavily were able to exploit resources retained in rhizomes from harvested culms. Nevertheless, small clumps produced much less seed than larger clumps, generating a risk of unknown magnitude that heavily harvested stands of monocarpic bamboos may be unable to satiate seed predators during their single opportunity for reproduction.     

EFFECTS OF AVIAN SEED DISPERSAL ON THE GENETIC STRUCTURE OF WHITEBARK PINE POPULATIONS

We used allozyme analysis to examine family structure, the spatial patterning of related individuals, in two populations of whitebark pine (Pinus albicaulis), a subalpine conifer that commonly displays a multistem form. The individual stems within clumps are genetically distinct individuals, having arisen from separate seeds. Individuals within a clump are genetically more similar than individuals in different clumps, but individuals in neighboring clumps do not appear to be more similar than individuals in distant clumps. This family structure appears to be a direct result of the seed-caching behavior of Clark's nutcrackers (Nucifraga columbiana), the primary dispersal agent for whitebark pine seeds.     

Changes in the Brown Seaweed Ascophyllum Nodosum (L.) Le Jol. Plant Morphology and Biomass Produced by Cutter Rake Harvests in Southern New Brunswick, Canada

Shoots and clumps of shoots of the commercial brown seaweed Ascophyllum nodosum (“rockweed”) add to the benthic complexity of the intertidal environment, providing an important habitat for invertebrates and vertebrates. To protect the structure of this habitat, management plans for the rockweed harvest of southern New Brunswick include restrictions on gear type and exploitation rates limited to 17% of the harvestable biomass. However, owing to physical and environmental factors, the harvest is not homogeneous, creating patches of exploitation ranging from 15 to 50%.A direct relationship existed between clump vulnerability, weight and length in a controlled harvest at 50% exploitation within 8 m by 8 m plots. At this exploitation rate, the gear rarely impacted clumps below 50 g or 60 cm in length. Clumps larger than 300 g and 130 cm were reduced by up to 55% of their length and 78% of their biomass. The overall impacts of the harvest on intertidal habitat is however of short duration as biomass recovers after a year of the experimental harvest. The rapid recovery is mostly due to a stimulation of growth and branching of the suppressed shoots of the clumps. Some harvested plots showed biomass even higher than initial levels, suggesting an increase in productivity at least during the first year after the harvest.     

Ecological benefits of integration of Calamagrostis epigejos ramets under field conditions

We established two independent experiments to estimate the ecological consequences of artificial severing on ramets of the competitively strong perennial grass Calamagrostis epigejos. We compared the responses of mature ramets of different size growing in different environments. Finally, we tested whether the response of young ramets to the severing depends on the density of surrounding vegetation.Severing decreased biomass and number of rhizomes of young ramets but did not affect their competitive tolerance. It decreased survival of mature ramets (probably due to traumata caused by cutting attached rhizomes) but did not influence total biomass of survived ramets. The response of total biomass of mature ramets to severing changed with size of the ramets. Further, biomass allocation to rhizomes changed differently after ramet severing in the two environments.The results suggest that field grown young ramets of Calamagrostis benefit from physiological integration. In contrast, mature ramets seem to be independent units according to the resource economy. Ecological benefits of integration depended on size of ramet clumps: ramets in clump had larger survival probability than control ramets. They also depended on environmental conditions: severing increased formation of new rhizomes at a sand dune subsrate, but it had a negative effect on plants in the forest experimental site. This intra-specific variation should be taken into account when trying to explain ecological patterns of integration benefits of clonal plants.     

Establishment of Salt Stress Tolerant Rice Plants Through Step Up NaCl Treatment In Vitro

Establishment of salt tolerant rice plants was examined by single step or step up NaCl treatments of shoot bud clumps in vitro, and variation among in vitro salt tolerant plants were examined by rapid amplified polymorphic DNA (RAPD). Shoot bud clumps were necrotic, stubbed or dead when subjected to single step treatment with 1.5 or 2.0 % NaCl. Conversely all the clumps could grow vigorously when subjected to step up salt treatment with 0.5, 1.0, 1.5 and 2.0 % NaCl at 3 week intervals and 2 % NaCl tolerant plants were established. RAPD revealed shoot bud clumps with and without different NaCl treatments, seedlings from field and grown in vitro, and regenerants from callus were genetically close to one another. Conversely, callus cultures were genetically isolated. Growth under different salt stress conditions was not correlated with the genetic variation, suggesting that 2.0 % NaCl tolerant plants might not result from genetic mutation but were due to adaptation of plants by step up NaCl treatment in vitro.     

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.     

Clonality in the Endangered <Emphasis Type="Italic">Ambrosia pumila</Emphasis> (Asteraceae) Inferred from RAPD Markers; Implications for Conservation and Management

Clonal plants have the ability to spread and survive over long periods of time by vegetative growth. For endangered species, the occurrence of clonality can have significant impacts on levels of genetic diversity, population structure, recruitment, and the implementation of appropriate conservation strategies. Here we␣examine clone structure in three populations of Ambrosia pumila (Nutt.) Gray (Asteraceae), a federally endangered clonal species from southern California. Ambrosia pumila is a perennial herbaceous species spreading from a rhizome, and is frequently found in dense patches of several hundred stems in a few square meters. The primary habitat for this species is upper terraces of rivers and drainages in areas that have been heavily impacted by anthropogenic disturbances and changing flood regimes. RAPD markers were employed to document the number and distribution of clones within multiple 0.25 m² plots from each of three populations. Thirty-one multi-locus genotypes were identified from the 201 stems sampled. The spatial distribution of clones was limited with no genotypes shared between plots or populations. Mean clone size was estimated at 9.10 ramets per genet. Genets in most plots were intermingled, conforming to a guerrilla growth form. The maximum genet spread was 0.59 m suggesting that genets can be larger than the sampled 0.25 m² plots. Spatial autocorrelation analysis found a lack of spatial genetic structure at short distances and significant structure at large distances within populations. Due to the occurrence of multiple genets within each population, the limited spread of genets, and a localized genetic structure, conservation activities should focus on the maintenance of multiple populations throughout the species range.     

Spatial distribution pattern of a clonal species: effects of differential production of clonal and sexual offspring

The spatial distribution patterns of genets and ramets within populations are expected to change as a function of the frequency with which clonal species recruit different types of offspring (sexual and clonal). We used an integrated approach to study the spatial arrangement of clonal plants by combining molecular and ecological data using Opuntia microdasys as a study system. The species is able to produce two types of clonal (plantlets and cladodes) and one type of sexual (seeds) offspring. Additionally it is found in three habitats that cause differences in the ability of each type of offspring to establish. In 2007, all individuals in the three habitats (162 in BH = bajada, 264 in IDH = hill-piedmont, and 136 at HPH = interdunes) were tagged and mapped. Amplified inter-simple sequence repeats (ISSR’s) were used to determine the multilocus genotype and relatedness of each individual ramet using 120 polymorphic bands (104 in BH, 128 in HPH and 180 in IDH). The spatial distribution pattern of genets and ramets was analyzed with the Hopkins test and spatial autocorrelation analysis. For all habitats we found that O. microdasys displayed a spatial distribution characterized by clumps of aggregated ramets, but habitats differed in the number of genets present. As for other clonal species a strong positive spatial autocorrelation exists within 20 m, although all analyses suggest that adjacent ramets are genetically less related to each other or belong to different genets, that is, ramets of different genets are intermingled. The spatial arrangement of genets and ramets in O. microdasys between habitats closely matches the frequency of establishment of each type of offspring (e.g. the more clonal areas are clumped groups of related individuals). These results confirm that in two habitats (BH and IDH) clonal recruitment had been more common than in the other habitat (HPH).     

Use of random amplified polymorphic DNA (RAPD) markers in the discrimination and verification of genotypes in Eucalyptus

We carried out four separate studies using random amplified polymorphic DNA (RAPD) markers to analyse samples of Eucalyptus supplied by several different organisations. The objective was to examine the reproducibility of the RAPD technique and its ability to discriminate between individual genotypes for verification of clonal identities. We found that RAPD profiles that are unique to a genotype can be generated reliably and simply and that even closely related genotypes can be distinguished. In addition, in each of the four studies, we detected cases where the plant material studied had been mis-sampled or mis-labelled (i.e. the RAPD profiles were not consistent with the identification numbers): (1) ramets of a Eucalyptus grandis clone were found to be derived from 2 different clones; (2) ramets labelled as 2 different Eucalyptus hybrid clones were found to be the same clone, owing to a mis-planted clonal hedge; (3) samples supplied as a single progeny of a controlled E. nitens cross were derived from two crosses involving different pairs of parents; (4) mis-labelling was detected for ramets of 4 of a set of 10 clones of E. grandis and E. camaldulensis. For three of the four studies, the detection of genotype mis-identifications was unexpected, suggesting that labelling or sampling errors during the handling of plant material are a frequent occurrence, with potentially serious economic consequences.     

Reproductive modes of three Ligularia weeds (Asteraceae) in grasslands in Qinghai-Tibet Plateau and their implications for grassland management

Ligularia virgaurea, L. sagitta and L. przewalskii are noxious weeds that are widely distributed in psychro-grasslands in the east of Qinghai-Tibet Plateau of China. However, little is known about reproductive modes and genetic diversity in the populations of L. virgaurea and L. sagitta under different environmental conditions, although such information is available for L. przewalskii. In this study, two experiments were conducted to examine the ex situ clonal growth of these weeds, and to determine the effects of reproductive modes on the diversity of their populations, using random amplified polymorphic DNA (RAPD) analysis. Transplanted seedlings of L. sagitta and L. virgaurea showed considerable clonal growth by forming rhizomes and ramets. The RAPD analysis showed that all individuals collected from natural populations were genetically different, indicating that no individuals originated from clonal growth. It is concluded that L. virgaurea and L. sagitta have the capacity for clonal growth, but the expression of their clonality is constrained by their growing conditions in natural grasslands. Sexual reproduction plays the major role in their population diversity in natural habitats, which requires that the control measures for these weeds should be targeted at the sexually reproducing populations.     

Microsatellite identification of ramet genotypes in a clonal plant with phalanx growth: The case of Cirsium rivulare (Asteraceae)

Cirsium rivulare is a perennial plant that forms patches consisting of ramets resulting from sexual reproduction by seeds and asexual propagation by rhizome fragmentation. We examined the relationship between the size of patches and genetic differentiation of ramets within and between patches. Ramet genotypes were identified using microsatellites. From among 216 ramets examined in the studied population, 123 had a unique genotype, while 93 were clonal, i.e., their genotype was present in at least two ramets. The frequency of ramets with clonal genotypes was 43% and the frequency of unique genotypes was 57%. Ramets with identical genotypes were dominant in small patches. Large patches consisted of ramets with both unique and clonal genotypes, usually with the predominance of the latter. A molecular variance analysis showed the highest level of variance between ramets and the lowest between patches. Additionally, 21.02% of the total variance was recorded between ramets and within patches. The size of patches was correlated with the number of clonal ramets and the number of unique ramets, but it was not correlated with the clonality index. This population of C. rivulare is currently in a phase of decline from 30 years of vegetation transformation, and there appears to have been an increase in sexual propagation based growth over clonal propagation based growth. Hence, a predominance of ramets with unique genotypes was observed. This can happen as a result of disintegration of large patches and formation of gaps between them. These gaps become convenient places for seed germination and the subsequent development of seedlings.     

Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest

The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy conditions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to different canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and cluster distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets’ biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F. nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. Moreover, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heterogeneous light environments. __________ Translated from Acta Ecologica Sinica, 2006, 26(12): 4019–4026 [译自: 生态学报]     

Spatially limited clonality and pollen and seed dispersal in a characteristic climber of Central African rain forests: Haumania danckelmaniana (Marantaceae)

Gene dispersal and clonality are important aspects of plant evolution affecting the spatial genetic structure (SGS) and the long‐term survival of species. In the tropics these parameters have mostly been investigated in trees and some herbs, but rarely in climbers which frequently: (1) show clonal growth leading to a patchy distribution pattern similar to that of understory herbs; and (2) flower in the canopy where they may have access to long‐distance dispersal like canopy trees. We thus hypothesize for climbers an intermediate genetic structure between herbs and trees. The study aims at assessing breeding system and spatial extent of clonality and gene dispersal in Haumania danckelmaniana (Marantaceae), a common climber in the tropical rain forests from western Central Africa. In eastern Cameroon, 330 ramets were sampled at three spatial scales and genotyped at seven microsatellite loci. Clonality was moderate (clonal extend: 15–25 m, clonal diversity 0.4–0.65) indicating the importance of recruitment from seeds at this locality. The low inbreeding (F_IS) suggested predominant outcrossing. The rate of decay of the relatedness between individuals with distance indicated limited gene dispersal distance (σ_g = 9–50 m, neighborhood sizes Nb = 23–67) in accordance with narrowly gravity dispersed seeds and restricted pollen transfer distance in densely flowering populations. The marked SGS (Sp = 0.011–0.026) was similar to that reported in tropical trees, but might increase with augmented clonality as in many herbs, especially under more severe disturbance regimes.     

Elevational patterns of species richness and density of rattan palms (Arecaceae: Calamoideae) in Central Sulawesi, Indonesia

We studied species richness and density of rattan palms in 50 plots of 10 × 100 m² each between 250 m and 2420 m in eight sites in Lore Lindu National Park (LLNP), Central Sulawesi. Rattans were observed in all sample sites, representing three genera and 34 species. The elevational patterns for species richness and density were humped-shaped with maxima around 1000 m. Polynomial models of second order explained 59 and 32% of species richness and density with the factor elevation, respectively. A majority of rattan species (65%) overlapped between 1000 and 1100 m elevation, while a pronounced change in the rattan flora occurred above 1100 m. Commercially important rattan species (Calamus zollingeri, C. ornatus var. celebicus, Daemonorops macroptera) were not observed above 1250 m. The change of species assemblage was significantly related to elevation (56%), followed by geographical distance (47%) and precipitation (40%). Less than 10% of LLNP is lowland forests, much of which is threatened by agricultural intensification. In contrast, montane forests are well represented in the park and high elevation forests are not subject to agricultural conversion or intensive harvesting of rattan and other forest products.     

Extensive clonality in the endangered shrub Haloragodendron lucasii (Haloragaceae) revealed by allozymes and RAPDs

The occurrence of clonality in threatened plants can have important implications for their conservation. In this study, allozymes and RAPDs were used to determine the extent of clonality in the endangered shrub Haloragodendron lucasii (Haloragaceae), which is known from only four sites within an 8 km range. Allozyme markers identified only six multilocus genotypes among the 53 ramets sampled across the four sites, although a total of 54 different genotypes were possible with the three polymorphic allozyme loci detected. The polymorphic bands detected in the RAPD analysis were capable of producing 2⁴⁶ genotypes, but again only six multilocus genotypes were delineated. The allozyme and RAPD data were congruent at three of the four sites. At the fourth site two genotypes were detected by each marker; however, once combined, three multilocus genotypes were observed. The probabilities that the observed number of replicates of each combined allozyme and RAPD genotype could be generated by sexual reproduction were less than 10^–18, leaving little doubt that clonality is the explanation for the observed patterns of genotypes. The genetic conclusions are supported by root excavations which show potential for vegetative reproduction and the observation of no sexual reproduction in the species. The recognition of extensive clonality in H. lucasii has had immediate implications for the conservation management of the species and resulted in changes to the management priorities for the species. Thus it is clear that appropriate genetic studies can play an important role in the management of threatened species.     

CLONE STRUCTURE IN FOUR SOLIDAGO ALTISSIMA (ASTERACEAE) POPULATIONS: RHIZOME CONNECTIONS WITHIN GENOTYPES

We studied clone structure and degree of genotypic mixing of Solidago altissima L. (Asteraceae) clones in four old fields near Ithaca, New York. The fields differed in time from agricultural abandonment and were approximately 1, 5, 20, and 35 years old. In the three older fields, three 0.75 m² plots were excavated intact and rhizome connections among ramets were mapped. In the youngest field 30 ramets were dug up singly. The genotype of all ramets was determined using electrophoresis of four polymorphic enzyme systems. Fields differed in the number and dispersion of genotypes within plots, and the degree of connection among ramets in the same clone. The one-year-old field was composed of single ramet genotypes which had probably established from seed the previous year. The five-year-old field contained many small contiguous clones of S. altissima with highly interconnected ramets. In the oldest two fields clones were highly intermixed and ramets of the same genotype were not extensively interconnected. These results demonstrate that clones of S. altissima display considerable phenotypic variability between fields and patterns of clone development may differ. The causes of this variability remain to be identified. We suggest that either selection for different genotypes or changing habitat conditions during succession may lead to changes in clone form.     

THE GENETIC CONSEQUENCES OF WORKER ANT POLLINATION IN A SELF-COMPATIBLE,CLONAL ORCHID

The self-compatible orchid Microtis parviflora is pollinated by the flightless worker caste of the ant Iridomyrmex gracilis. The orchid is clonal and forms small patches, usually less than 1 m², of disconnected individual ramets. Ant pollinators visited and revisited a limited proportion of available inflorescences, and 40% of all flower visits occurred within plants promoting self-pollination. Pollen labels indicated that self-pollination accounted for 51% of the pollen transfers, although pollen carryover extended beyond 16 flowers on 2 or 3 inflorescences. The distribution of ant movements between plants was leptokurtic with a mean of 12.4 ± 14.9 cm and a maximum of 89 cm, but a high proportion of movements were within clones accentuating the level of self-pollination. However, some pollen transfers between inflorescences of unlike genotypes contributed to a low incidence (max = 8%) of outcrossing. In 12 patches examined by electrophoresis, the density varied from 11 to 61 inflorescences per m² and a maximum of only 4 genotypes were detected. Electrophoretic analysis revealed populations were highly inbred: only 23% (N = 17) of the loci were polymorphic and the mean gene diversity h, was 2.7%. Heterozygotes were observed in only one population given a mean fixation index F, of 0.982. These results reflect the combined effects of restricted ant foraging and clonality. Nevertheless, while ant foraging was restricted, some outcrossing occurred and in the absence of clonality it is likely that ant foraging would have yielded a mixed mating system similar to those reported for a wide array of insect pollinators. Given the ability of ants to generate pollen flow, the reasons for the rarity of ant pollination appear to lie elsewhere.     

SPATIAL DISTRIBUTION OF GENETIC INDIVIDUALS IN THICKETS OF ALNUS INCANA SSP. RUGOSA,A CLONAL SHRUB

Thickets of speckled alder (Alnus incana ssp. rugosa (Du Roi) Clausen) consist of numerous discrete clumps of stems. Presumably all stems in a single clump are part of a single genetic individual, but a genet could comprise more than one clump. Starch-gel electrophoresis was used to identify genetic individuals in four alder populations in central New York. A single genetic marker, a tetramer with three alleles, could discriminate five genotypes. Nearest neighbor analysis revealed that genotypes were distributed randomly. That is, the pattern of genotypes was statistically indistinguishable from a model where each clump is considered a unique individual and where clump genotypes are randomly distributed. Calculation of Morisita's index of dispersion confirmed that clumps of a single genotype were not aggregated. Although alder is capable of forming root suckers and offsets, lateral expansion of genets is apparently ineffective. Apparently, spatial distribution of genetic individuals within alder thickets is not influenced by clonal growth or by other factors acting to cause patterns in the genetic structure of plant populations.     

Analysis of the multi-stem clump structure ofLitsea japonica Juss. growing in a coastal dwarf forest

The multi-stem clump structure of a coastal dwarf forest dominated byLitsea japonica Juss. was investigated in order to clarify the sprouting characteristics and self-maintenance of clumps by stem alternation. The size and age distribution of multi-stem clumps were analyzed using cumulative relative frequency curves.L. japonica had a large number of stems and an even height distribution or young age-biased distribution of stems within a clump. These results indicated the sequential flushing of sprouts at high frequency. Height distribution within a clump ofL. japonica was relatively even compared to other species. This clump structure suggested the stable self-maintenance of individuals in all ranges of size and age without disturbances. It originated specific sprouting characteristics as a response to the severe stress of salty wind.Ardisia sieboldii Miq. had few stems within a clump. Although the stem height distribution of large individuals tended to be even, most clumps had a large size-biased distribution of stem height which indicated simultaneous sprouting. From this structure, sprouts of this species were thought to be of less significance in the stable self-maintenance of individuals thanL. Japonica.