Variation in growth rate in shoot populations of the clonal dwarf shrub Linnaea borealis

This study was based on two years of field observations of a Linnaea borealis population in a coniferous forest in Sweden. Detailed information on performance and survivorship of shoots were gathered from consecutively made drawings of 90 individually marked shoot systems of Linnaea distributed among 15 plots in the forest. Branching of the shoot systems was correlated to growth of the main shoot. Increasing light flux negatively affected growth of Linnaea . A tendency for a competitive effect on Linnaea of Vaccinium spp. was also found. In contrast, the surrounding mosses, and intraspecific density of Linnaea had no significant effect on the shoot systems. The growth rate of the shoot population was analysed by matrix models. Population growth rate was most dependent on survivorship and growth of main shoots in Linnaea , while the survivorship of lateral shoots was of less importance. In spite of an average decrease in the population size as revealed by the matrix model simulation, the incorporation of the observed spatial variation in shoot growth increment, branching and shoot survivorship, yielded varying population growth rates. Parts of the population increased in size. The importance of choice of spatial scale in population analyses of forest understory plants is emphasized.     

Stochastic population dynamics of clonal plants: Numerical experiments with ramet and genet models

Dynamics of ramer and genet populations were analyzed by use of stochastic matrix models. Based on field data, population development and extinction rates during 50 simulated years were estimated for ramet populations of three speciesPotentilla anserina, Rubus saxatilis andLinnaea borealis. Only small initial populations (below 125–250 ramets), experienced a detectable risk of extinction within this time interval. ForP. anserina andR. saxatilis, population increase occurred in some simulations despite negative average growth rates. A model for stochastic genet dynamics was constructed by combining field data and hypothesized parameter values. Growth rate and population structure were insensitive to variation in disturbance intensity and frequency, whereas variation in recruitment affected population structure but only to a minor extent growth rate. Decreasing recruitment causes extinction of genet populations, but the time-scale for the decline is in the magnitude of centuries for initial genet populations of about 1000 individuals. Dynamics of genets in clonal plants thus incorporate processes occurring on widely different scales. Some implications of the results for models of population dynamics in long-lived clonal plants are discussed.     

Sexual reproduction, clonal diversity and genetic differentiation in patchily distributed populations of the temperate forest herb Paris quadrifolia (Trilliaceae)

Clonal plant species have been shown to adopt different strategies to persist in heterogeneous environments by changing relative investments in sexual reproduction and clonal propagation. As a result, clonal diversity and genetic variation may be different along environmental gradients. We examined the regional and local population structure of the clonal rhizomatous forest herb Paris quadrifolia in a complex of forest fragments in Voeren (Belgium). Relationships between population size (the number of shoots), shoot density (the number of shoots per m²) and local growth conditions were investigated for 47 populations. Clonal diversity and genetic variation within and among 19 populations were investigated using amplified fragment length polymorphism markers. To assess the importance of sexual reproduction, seed set, seed weight and germination success were determined in 18 populations. As predicted, local growth conditions largely affected population distribution, size and density of P. quadrifolia. Populations occurring in moist and relatively productive sites contained significantly more shoots. Here, shoots were also much more sparsely distributed compared to populations occurring in dry and relatively unproductive sites, where shoots showed a strongly aggregated distribution pattern. Clonal diversity was relatively high, compared with other clonal species (G/N ratio = 0.43 and Simpson’s D=0.81). Clonal diversity significantly (P<0.01) decreased with increasing shoot density while molecular genetic variation was significantly (P<0.01) affected by population size and local environmental conditions. Lack of recruitment and out-competition of less-adapted genotypes may explain the decreased genetic variation in dry sites. Analysis of molecular variance revealed significant genetic variation among populations (Φ _ST=0.42, P<0.001), whereas pairwise genetic distances were not correlated to geographic distances, suggesting that gene flow among populations is limited. Finally, the number of generative shoots, the number of seeds per fruit and seed weight were significantly and positively related to population size and local growth conditions. We conclude that under stressful conditions populations of clonal forest plant species can slowly evolve into remnant populations characterized by low levels of genetic variation and limited sexual reproduction. Conservation of suitable habitat conditions is therefore a prerequisite for effective long-term conservation of clonal forest plant species.     

Shoot growth dynamics and size-dependent shoot fate of a clonal plant,Festuca rubra, in a mountain grassland

The relation of the within-season and between-season patterns of shoot growth were compared in a clonal grass with long-lived shoots,Festuca rubra, in a mown mountain grassland. The growth rate of shoot length from spring to summer in a year was almost constant for each shoot irrespective of spring shoot length each year. The annual shoot growth rate from spring to spring was negatively correlated with the shoot length in the first spring. Shoots of different length and age therefore tended to converge over time to a population of identical shoot size, suggesting an equalizing effect of growth pattern on size structure. Shoot size (shoot length and number of leaves) influenced the fates of shoots. Larger shoots showed an increased incidence of both flowering and formation of intravaginal daughter shoots and a decreased incidence of death in the subsequent time period. The fates of shoots were independent of their age. Although the negatively size-dependent springto-spring annual shoot growth rate acted to decrease shoot size variation, the remaining variation within the shoot population was still sufficient to generate different fates of shoots. These fates were not related to the previous life history of individual shoots. There was a significantly positive effect of the shoot size at initiation on its life expectancy. This was mainly attributable to the positively size-dependent survival rate of shoots in the early stage (<1 year old) of shoot life history. Later on (> 1 year old), shoot size had little effect on the survival rate of shoots. Once small young shoots have survived this early stage (< 1 year old) in life history, they can grow vigorously, little affected by competition regardless of shoot size, and converge to a stable size structure of shoots of similar size. Only shoot size in the early stage ( < 1 year old) of life history is important for the persistence of a shoot population.     

Estimation of dieback process caused by herbivory in an endangered root-sprouting shrub species,Paliurus ramosissimus (Lour.) Poir., using a shoot-dynamics matrix model

Mortality, a critical parameter for population dynamics, is difficult to measure in long-lived trees or clonal herbaceous species because of the extremely low frequency of deaths. A model based on shoot recruitment would be helpful to estimate the population fate of species without a sufficient number of observed deaths. Existing matrix models are applicable to the dynamics of physiologically independent shoots, but not to physiologically dependent ones. We developed a shoot-dynamics model for plants with physiologically-dependent shoots, and used the model to estimate the effects of herbivory and conservation measures on the dynamics of a long-lived, shoot-sprouting shrub species, Paliurus ramosissimus (Rhamnaceae). Two populations of the endangered shrub have been severely damaged by herbivory by sika deer. The damage was mainly to new sprouting shoots. No deaths of individual plants were observed during an 8-year survey, and we could not estimate mortality. Thus, prediction of population dynamics based on births and deaths of individuals was impossible. Because P. ramosissimus is a shoot-sprouting species, we instead estimated the decline of individuals using a shoot-dynamics model. Using this model, we estimated the time to an 80 % decrease in shoot number per individual in the two populations at 37.8 and 37.2 years. These lengths suggest an immediate need for conservation measures to prevent herbivory even though no death of any individual was observed in the field survey. The estimated recovery times from the herbivory damage were 28.7 and 29.2 years if herbivory of new shoots is completely prevented by conservation measure.     

Growth and competition in clonal plants-persistence of shoot populations and species diversity

This paper reviews studies on growth and size-structure dynamics of shoots and clones in clonal plants in comparison with those in non-clonal plants, and discusses the characteristics of clonal plants. The mode of competition between individuals (symmetric versus asymmetric, degree of competitive asymmetry), growth dynamics of individuals, allocation pattern between organs and spatial pattern of individuals are closely correlated with each other in non-clonal plant populations. Theoretical and field studies based on the diffusion model revealed that plants of “height-growth” type (mostly early-successional tree species) and plants of “diameter-growth” type (mostly late-successional tree species) tend to exhibit asymmetric competition and symmetric competition respectively. Moreover, asymmetrically competing plants show smaller effects of variation in individual growth rate and spatial pattern on the size-structure dynamics of the population than symmetrically competing plants. Thefefore, the spatial pattern of inviduals should be considered especially for plants undergoing symmetric competition. These results for non-clonal plants should have a significant implication also for the growth dynamics and competition in clonal plants. The mean growth rate of shoots [G(t,x) function] and hence the mode of competition between shoots differs among clonal plant species as in non-clonal plants. However, a large magnitude and size-independence (or slightly negative size-dependence) of the variation in growth rate of shoots [D(t,x) function], especially at the early stage in a growing season is a common characteristic of many clonal plant species, in contrast to the positively size-dependent variation in individual growth rate in non-clonal plants. This type of variation in shoot growth rate leads to the persistence of stable shoot populations even when the mean growth rate function is changed, and also in cases where the shoot population structure would be unstable in the absence of variation in growth rate. It is suggested that competition between clones is symmetric in most clonal plant species, which brings about small-scale spatio-temporal changes in species abundance and hence species diversity.     

Competition between Shoots in Stands of Clonal Plants

Abstract Data on shoot turnover, shoot size distributions and individual shoot growth in stands of clonal plants are reviewed and compared with shoot density regulation in non-clonal plants. Shoots are the smallest units capable of independent existence in clonal as well as unitary plants and this comparison allows an investigation of the impact of integration and other factors on clonal shoot dynamics. Physiological integration, the capacity of clonal plants to exchange resources and growth substances between interconnected shoots, has long been held responsible for the ability of clonal plants to prevent shoot overproduction and for the general lack of density-dependent mortality and self-thinning. A review of published data leads to the conclusion that the effects of integration on shoot dynamics in clonal plants have been overestimated. Other characteristics such as the density-dependent control of meristem activity, the synchronization of shoot growth in spring and the maximum shoot size, seem more important for the deviating behaviour of shoot populations of clonal plants.     

Longevity of clonal plants: why it matters and how to measure it

Background

Species'' life-history and population dynamics are strongly shaped by the longevity of individuals, but life span is one of the least accessible demographic traits, particularly in clonal plants. Continuous vegetative reproduction of genets enables persistence despite low or no sexual reproduction, affecting genet turnover rates and population stability. Therefore, the longevity of clonal plants is of considerable biological interest, but remains relatively poorly known.

Scope

Here, we critically review the present knowledge on the longevity of clonal plants and discuss its importance for population persistence. Direct life-span measurements such as growth-ring analysis in woody plants are relatively easy to take, although, for many clonal plants, these methods are not adequate due to the variable growth pattern of ramets and difficult genet identification. Recently, indirect methods have been introduced in which genet size and annual shoot increments are used to estimate genet age. These methods, often based on molecular techniques, allow the investigation of genet size and age structure of whole populations, a crucial issue for understanding their viability and persistence. However, indirect estimates of clonal longevity are impeded because the process of ageing in clonal plants is still poorly understood and because their size and age are not always well correlated. Alternative estimators for genet life span such as somatic mutations have recently been suggested.

Conclusions

Empirical knowledge on the longevity of clonal species has increased considerably in the last few years. Maximum age estimates are an indicator of population persistence, but are not sufficient to evaluate turnover rates and the ability of long-lived clonal plants to enhance community stability and ecosystem resilience. In order to understand the dynamics of populations it will be necessary to measure genet size and age structure, not only life spans of single individuals, and to use such data for modelling of genet dynamics.     

Current-year shoot based approach for annual variation in the reproductive output in Siberian alder (<Emphasis Type="Italic">Alnus hirsuta</Emphasis> var. <Emphasis Type="Italic">sibirica</Emphasis>)

Siberian alder (Alnus hirsuta var. sibirica) shows annual variation in reproductive output. However, this phenomenon has not been explained by hypotheses proposed in previous studies. In this study, we constructed a matrix model of current-year shoot dynamics to estimate and compare the reproductive output of current-year shoot population with or without annual variation, and explained the annual variation of reproductive output in terms of a reproductive strategy of the current-year shoot population of Siberian alder. The current-year shoot population of Siberian alder was divided into three functional groups: reproductive, maintenance, and explorative shoot sub-populations. A transition matrix was calculated from the relationships between 1-year-old shoots and the current-year shoots on them. The dynamics of the current-year shoot population was simulated using the estimated matrix on the patterns with or without annual variation in reproductive output. The pattern with annual variation in reproductive output yielded more reproductive current-year shoots than the pattern with high reproductive output every year. The annual variation of reproductive output may well be regarded as a reproductive strategy of current-year shoots to increase lifetime fecundity.     

Demographic population structure of a leafy epixylic hepatic Anastrophyllum hellerianum (Nees ex Lindenb.) R.M.Schust

Understanding the population processes is central when assessing proper conservation acts, especially in the case of species which inhabit temporal and patchy substrates, such as decaying wood. In epixylic bryophytes, however, only little is known of the demographic population structure and within-population dynamics. In this study on a leafy epixylic hepatic, Anastrophyllum hellerianum (Nees ex Lindenb.) R.M.Schust., a set of demographic traits is investigated, including colony area and size, proportions of the sterile, gemmiparous or sexual developmental stages, the shoot size class distribution among the developmental stages, and numbers of asexual propagules present on the shoots. The estimates of mortality are assessed on the basis of the proportions of dead shoots found in the colonies. The size distribution within the developmental stages of this species shows that a certain threshold size is required for both asexual and sexual reproduction. The shoot size is negatively correlated with density, whereas the numbers of asexual gemmae and the proportions of developmental stages are not affected by shoot density. Furthermore, provisional estimates are calculated for the transitions from early developmental stages to reproductively mature stages. The results indicate that the initial phases of the gametophyte development are the most vulnerable.     

Translocation of twinflower (Linnaea borealis L.) in the Scottish Borders

Summary

Twinflower, Linnaea borealis L., is a creeping, woody, clonal perennial nationally scarce in the UK, with its distribution confined to discrete patches in Scotland. Translocation of twinflower from natural patches into either existing patches, with a view to increasing genetic variation and outcrossing rates, or to unoccupied habitat, with a view to increasing its overall prevalence, could effectively increase the species' viability. In a small pilot experiment 38 shoots were moved from a vigorous patch in the Scottish Borders into two overtly hospitable clearings within the same woodland. Two years later 18 shoots (47%) survived and three years later four shoots (10.5%) remained, with die-back occurring irregularly over shoots of different initial lengths. No measured character of the original shoots explained which survived at each time interval. The most likely cause of the high mortality was competition from established grasses (Deschampsia flexuosa) which prevented growth and the formation of new roots along stolons. The study should help focus future efforts to determine best methods of increasing establishment of twinflower in Scotland.     

Shoot demography in new England populations of Maianthemum canadense desf.

Summary The demography of shoots of eight populations of a herbaceous perennial exhibiting clonal growth, is presented. The study was done along an elevational gradient, from a more open secondary mixed forest to a denser, more mature stand. Most shoots lived one to three years on the average, but shoots as old as twelve years were found. Large variation in formation and mortality of shoots was observed among plots and years. Yearly trends in the mortality rates of site replicates showed a higher correlation than rates of shoot formation. Although the density of shoots was highest in the drier sites, the turnover of shoots was highly variable and apparently uncorrelated with site location. Age structures revealed a tendency of longer-lived and higher reproductive activities among shoots from more mesic sites. It is hypothesized that environmental rather than density controls are primary causes of the population dynamics observed in this species.     

Demographic vulnerability of the clonal and endangered meadow thistle

For effective management of endangered species it is pivotal to understand why a species is endangered and which key life cycle components are involved in its response to environmental changes. Our objective was to investigate the response of rosettes of the redlisted clonal herb Cirsium dissectum to anthropogenic nutrient enrichment, which threatens its populations, and the consequences of these responses for its population dynamics. We constructed matrix population models with demographic data from three populations and four annual transitions and we decomposed the spatiotemporal variation in projected population growth rates into contributions from life cycle components. These patterns were compared with below-ground rosette dynamics in different fields, and with the below- and above-ground rosette dynamics in a garden experiment with nutrient enrichment and competing grasses. The decomposition analysis revealed that increased clonal rosette formation and decreased rosette survival were driving the spatial variation in the population growth rate. Excavating the below-ground rhizome network revealed a higher rosette turn-over in experimentally fertilized garden plots, which not only resulted in increased plot-level extinction, but also in increased spread of the clonal offspring. This supported the observed trend among field populations: rosette formation trades off with rosette survival. Surviving seedlings were only found in areas where the topsoil had been removed. The endangered C. dissectum is vulnerable when its habitat becomes more productive, because this species does not have the necessary capability to build up biomass. Small-scale disturbances such as created by sod-cutting or trampling cattle are essential for seedling establishment and necessary to render the explorative strategy of rhizomatous clonal spread successful.     

Shoot dynamics of the giant grass Gynerium sagittatum in Peruvian Amazon floodplains,a clonal plant that does show self-thinning

The giant rhizomatous grass Gynerium sagittatum is an early successional species that forms dense monocultures in Peruvian Amazon floodplains. We studied the shoot population structures by recording shoot densities and shoot heights. Leaf areas and stem volumes were allometrically estimated. Stands of two varieties of G. sagittatum were examined that differ in height and in the degree of shoot branching. In stands of increasing age, marked decreases in shoot densities were accompanied with an increase in mean shoot size. Self-thinning was indicated by the negative correlation between log stem volume per unit ground area and log shoot density, significant at least for one of the two varieties. The difference in thinning slope between the varieties could be largely accounted for by their different shoot geometry, as was revealed by calculations based on the allometric model of Weller (1987b). The relationship between log leaf area per shoot and log shoot density was significantly negative with slopes close to –1. Shoot size inequalities decreased with increasing mean stem volume per shoot, probably as a result of density-dependent mortality of the smaller shoots. All of these results accord with expectations for shoot self-thining. Gynerium sagittatum is the first clear example of a clonal plant species that exhibits self-thining in natural monospecific stands. It is argued that self-thinning occurs in this giant tropical grass because its shoots are perennial and do not experience seasonal die-back (periodic density-independent mortality), in contrast to many of the clonal plant species that have been studies so far.     

Microgeographic variation in recruitment under adult trees: arrival of new genotypes or perpetuation of the existing ones?

• Investigating spatial variation in the relative importance of sexual reproduction and clonal propagation is critical to obtain more accurate estimates of future effective population sizes and genetic diversity, as well as to identify ecological correlates of clonality.
• We combined a stratified sampling scheme with microsatellite genetic analyses to estimate variation in the proportion of sexual versus clonal recruits among saplings in five populations of the tree Pyrus bourgaeana. Using a likelihood framework, we identified clones among the genotypes analysed and examined variation among populations regarding the proportion of saplings coming from clonal propagation. We also examined the relationship between the relative abundance of clonal shoots across the studied populations and their herbivory levels.
• Our results revealed that one third of the saplings examined (N = 225 saplings) had a probability above 0.9 of being clones of nearby (<10 m) trees, with the ratio between clonal propagation and sexual recruitment varying up to eight‐fold among populations. A small portion of these putative clonal shoots reached sexual maturity. Relative abundance of clonal shoots did not significantly relate to the herbivory by ungulates.
• Our results call into question optimistic expectations of previous studies reporting sufficient levels of recruitment under parental trees without animal seed dispersal services. Nevertheless, given that some of these clonal shoots reach sexual maturity, clonal propagation can ultimately facilitate the long‐term persistence of populations during adverse periods (e.g. environmental stress, impoverished pollinator communities, seed dispersal limitation).

     

Effect of reproductive modes and environmental heterogeneity in the population dynamics of a geographically widespread clonal desert cactus

The dynamics of plant populations in arid environments are largely affected by the unpredictable environmental conditions and are fine-tuned by biotic factors, such as modes of recruitment. A single species must cope with both spatial and temporal heterogeneity that trigger pulses of sexual and clonal establishment throughout its distributional range. We studied two populations of the clonal, purple prickly pear cactus, Opuntia macrocentra, in order to contrast the factors responsible for the population dynamics of a common, widely distributed species. The study sites were located in protected areas that correspond to extreme latitudinal locations for this species within the Chihuahuan Desert. We studied both populations for four consecutive years and determined the demographic consequences of environmental variability and the mode of reproduction using matrix population models, life table response experiments (LTREs), and loop and perturbation analyses. Although both populations seemed fairly stable (population growth rate, λ∼1), different demographic parameters and different life cycle routes were responsible for this stability in each population. In the southernmost population (MBR) LTRE and loop and elasticity analyses showed that stasis is the demographic process with the highest contributions to λ, followed by sexual reproduction, and clonal propagation contributed the least. The northern population (CR) had both higher elasticities and larger contributions of stasis, followed by clonal propagation and sexual recruitment. Loop analysis also showed that individuals in CR have more paths to complete a life cycle than those in MBR. As a consequence, each population differed in life history traits (e.g., size class structure, size at sexual maturity, and reproductive value). Numerical perturbation analyses showed a small effect of the seed bank on the λ of both populations, while the transition from seeds to seedlings had an important effect mainly in the northern population. Clonal propagation (higher survival and higher contributions to vital rates) seems to be more important for maintaining populations over long time periods than sexual reproduction.     

Structure and Dynamics of Populations of Japanese Barberry (Berberis Thunbergii DC.) in Deciduous Forests of New Jersey

Japanese barberry, Berberis thunbergii DC., has become a prominent exotic species in deciduous forests throughout the eastern and midwestern US. Populations range from small plants occurring at low densities to dense, impenetrable thickets of plants with up to 40 stems/individual. A study was undertaken at Morristown National Historical Park in New Jersey to document plant densities, plant size, recruitment through vegetative processes of new shoot initiation and clonal spread and recruitment from seedling establishment, and mortality of stems and plants. Nearly 2000 shoots on 370 plants were individually marked and followed for two growing seasons, and over 1000 seedlings were also individually marked and followed. Populations vary much more in total shoots/area than they do in plant individuals/area, or in mean plant size (shoots/plant), as even the sparse populations have a few large individuals. Shoot mortality is less than new shoot initiation, but most plants do not change in size or change by small numbers of stems. However, the number of new shoots per plant increases as plant size increases. Once plants have three stems, they suffer little or no mortality. Seedling establishment is proportional to the density of shoots, so that as plants grow in size, local recruitment from seed increases. Large numbers of seedlings, and a survival rate of 10%, combine to make seedling recruitment a major component of population increase. The combination of multiple forms of vegetative and seed-based population growth, and the very low rates of plant mortality due to the multi-stemmed growth form explains the ability of this invasive species to rapidly produce dense, persistent populations.     

慈竹母株大小对克隆生长的影响

 克隆生长在慈竹种群更新中占有重要地位。本文以基径和生物量两个指标同时衡量慈竹(Neosinocalamus affinis)的母株大小,并分析了母株大小对新竹笋产生、竹笋大小和根茎大小的影响。结果表明：1）慈竹的成竹基径与一级母株基径、二级母株基径以及与2龄和3龄母株平均单株生物量呈正相关,但是成竹基径随一级母株基径增加比随二级母株基径增加而增加的速率快得多;2）慈竹母株大小与各样方中出笋数、活笋数和死笋数无显著的正相关;3）发笋母株的基径显著大于未发笋母株,但发一个笋母株与发两个以上笋母株的基径无显著差异;4）母株基径、2龄和3龄母株平均单株生物量都与根茎的长度和粗度呈显著正相关,且竹笋基径与根茎粗度也呈显著的正相关。母株大小对慈竹竹笋的影响是通过影响根茎实现的。总之,慈竹的克隆生长在个体水平受母株大小制约。     

Clonal distribution,fertility and sex ratios of the moss Plagiomnium affine (Bland.) T.Kop. in forests of contrasting age

Abstract

Six populations of the clonal forest floor moss Plagiomnium affine from forests of different age were screened for genetic variation at 23 allozyme loci, of which nine were polymorphic. Samples consisting of two adjacent unconnected shoots were taken at regular intervals along one transect from each population. A total of 602 shoots was analysed. Almost 80% of the shoots were sterile (i.e. not expressing male or female gender). Sex remained unknown for only 10% of shoots after identification of genets based on electrophoretic data. We identified a mean number of 3.7 fertile clones per population. The mean length of clones along transects in each population ranged between 2 and 3 m. The size distribution within populations was bimodal, with a few dominant clones and a varying number of much smaller clones. The overall sex ratio was slightly female biased at the ramet level, but balanced at the genet level. Forest age was negatively correlated with percentage of sterile shoots and positively correlated with frequency of sporophytes. In both cases correlations were significant only if population 1, which was subject to extreme soil disturbance by badgers, was excluded. We conclude that the effective population size is larger, and the susceptibility to genetic drift is lower, in old forests.     

Local forest environment largely affects below-ground growth, clonal diversity and fine-scale spatial genetic structure in the temperate deciduous forest herb Paris quadrifolia

Paris quadrifolia (herb Paris) is a long-lived, clonal woodland herb that shows strong differences in local population size and shoot density along an environmental gradient of soil and light conditions. This environmentally based structuring may be mediated by differences in clonal growth and seedling recruitment through sexual reproduction. To study the interrelationship between environmental conditions and spatial patterns of clonal growth, the spatial genetic structure of four P. quadrifolia populations growing in strongly contrasting sites was determined. In the first place, plant excavations were performed in order to (i) determine differences in below-ground growth of genets, (ii) investigate connectedness of ramets and (iii) determine total genet size. Although no differences in internode length were found among sites, clones in moist sites were much smaller (genets usually consisted of 1-3 interconnected shoots, most of them flowering) than genets in dry sites, which consisted of up to 15 interconnected shoots, the majority of which were vegetative. Further, amplified fragment length polymorphism (AFLP) markers were used. Clonal diversity was higher in populations located in moist and productive ash-poplar forests compared to those found in drier and less productive mixed forest sites (G/N: 0.27 and 0.14 and Simpson's D: 0.84 and 0.75, respectively). Patterns of spatial population genetic structure under dry conditions revealed several large clones dominating the entire population, whereas in moist sites many small genets were observed. Nevertheless, strong spatial genetic structure of the genet population was observed. Our results clearly demonstrate that patterns of clonal diversity and growth form of P. quadrifolia differ among environments. Limited seedling recruitment and large clone sizes due to higher connectedness of ramets explain the low clonal diversity in dry sites. In moist sites, higher levels of clonal diversity and small clone sizes indicate repeated seedling recruitment, whereas strong spatial genetic structure suggests limited seed dispersal within populations.