Population structure and dynamics of the clonal dwarf-shrub Linnaea borealis

Abstract. The demography of the long-lived clonal dwarf-shrub Linnaea borealis was studied during four years in a coniferous forest in central Sweden. The main object was to infer patterns of temporal variation in population dynamics of this species. The shoot population is organized in fragments, i.e. physically connected systems of shoots partly covered by the moss carpet. The age and size structure of the fragment population is described, but shoots are more convenient units for a study of population dynamics. A stochastic model of shoot population dynamics was constructed, and simulations indicated a considerable temporal variation in population size and flowering. Hence, variability as such is an essential aspect of the dynamics of established populations of Linnaea. Simulations of extinction risks revealed that small-sized shoot populations (ca. 250 shoots) are likely to be long-lived when experiencing environmentally induced demographic variation of the range observed. Mortality agents for established genets, such as large-scale disturbances, were not incorporated in the models. Some implications of variable population growth rates in clonal plants in woodlands are discussed.     

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.     

Ramet size equalisation in a clonal plant,Phragmites australis

The influence of shading from older generations of dead culms (standing litter) on density, growth rate and development of size structure at the ramet level was investigated in a pure stand of Phragmites australis by experimental neutral shading of plots after removal of standing litter. Initial differences in height distribution between autumn and spring cohorts disappeared in the course of shoot growth. The Gini coefficients of shoot heights and estimated shoot weights indicated that the size structure of the shoots became more equal with increasing mean size in both shaded and unshaded plots. Relative growth rate for height (RHGR) and weight of individual shoots was negatively related to shoot size during the early and presumably storage-dependent growth period, suggesting a strong support for growth of smaller shoots. No etiolation was indicated by mean or maximum height in shaded and unshaded plots, or by the relationship between shoot height and weight. Mean shoot density was significantly lower in shaded than in unshaded plots in one of two shade treatment years. A regression model indicated a small but significant effect of shoot density on the approximately linear relationship between RHGR and the logarithm of height. The growth rate of small shoots was slightly larger at low than at high shoot density. Therefore, it is suggested that the shade from standing litter in P. australis stands can decrease shoot natality in the spring cohort, and thereby increase the support to fewer small shoots.     

Population growth of a gorgonian coral: equilibrium and non-equilibrium sensitivity to changes in life history variables

Summary A size dependent model of population growth of the Caribbean gorgonian Plexaura A is developed based on observed rates of survival, growth and colony fragmentation at a site in the San Blas Islands, Panama. Sensitivity and elasticity analyses indicate that the fate of large colonies has the greatest effect on population growth. Variables which directly affect the generation of large colonies have the next greatest effect on population growth. These variables include the recruitment of large fragments, and the survivorship of colonies in the next smaller size class. Sexual reproduction has an extremely limited ability to affect population growth. Vegetative reproduction has a greater potential effect on growth rates. Environmental conditions regularly change the matrix of transition probabilities which predicts population growth. This keeps the population from approaching its stable size class distribution. Deviations from the stable size class distribution alter sensitivity and elasticity and in this case have the effect of increasing the importance of survivorship of the smallest colonies. Nonequilibrium conditions alter sensitivity analyses and it is important to assess whether populations are at equilibrium and to determine the effects of such deviations on the sensitivity analysis.     

Shoot life span in relation to successional status in deciduous broad-leaved tree species in a temperate forest

In trees, leaf life span is closely related to successional status. Although leaves are attached to shoots, shoot life span has been insufficiently studied in the context of ecological systems. Interspecific variation in shoot survivorship was investigated over 27 months in 15 temperate hardwood tree species. Relationships between shoot architecture and shoot survival were also investigated. Shoot life span was shortest in early successional species, and longest in late successional species, in each of the families Betulaceae and Fagaceae. In Salicaceae, all of which were early successional species, shoot life span was longer in mountainous than in riparian species. Early successional or riparian species distributed longer shoots densely, even in proximal positions on mother shoots, resulting in mutual shading and consequent early and massive shoot shedding. By contrast, late successional or mountainous species concentrated shoots in distal positions, allowing shoots to receive equally favorable light, resulting in a longer life span. These results reveal close relationships between shoot life span and environmental resource availability or successional status and suggest a causal relationship between shoot shedding and shoot architecture.     

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.     

Growth and development of Potamogeton distinctus in an irrigation pond in SW Japan

Growth and development of a population of Potamogeton distinctus were studied over one vegetation period.
The morphological structure of the species is described. Maximum development of the size of shoot complexes occurred in August (up to 12 g dry weight/shoot complex). The plant exhibits a regular growth cycle with continuous horizontal and vertical growth that is illustrated by the development of horizontal shoots, vertical shoots and different plant parts. The lower horizontal shoots allow an extension up to 1.8 m from the point of germination. Before the rainy season in June the 1st to 5th generations of vertical shoots mainly contribute to the standing crop of the plants' stand, and after the rainy season the 5th to 9th generation shoots are predominant. Floating leaves make up the main proportion of biomass during the whole growth cycle. Almost 100% of the shoot complexes were found flowering and producing numerous seeds. Up to 16% of the standing crop are allocated to inflorescence and seeds in August. Turions failed to form that year because of rapid desiccation of the habitat. The observed average lifetime of a vertical shoot is appr. 65 days, the estimated turnover rate is between 2.15 and 2.27.     

Relationship between reproductive behavior and new shoot development in 5-year-old branches of olive trees (Olea europaea L.)

The development of new shoots plays a central role in the complex interactions determining vegetative and reproductive growth in woody plants. To explore this role we evaluated the new shoots in the olive tree, Olea europaea L., and the effect of fruiting on new shoot growth and subsequent flowering. Five-year-old branches served as canopy subunits in order to obtain a global, whole-tree view of new shoot number, size and morphological origin. The non-bearing trees had many more shoots than the fruit-bearing trees, and a greater number of longer shoots. In both bearing conditions, however, the majority of shoots were less than 4 cm long, with shoots of progressively longer lengths present in successively decreasing frequencies. Six major shoot types were defined on the basis of apical or lateral bud origin and of parent shoot age. On fruit-bearing trees, the new shoots originated predominantly from the shoot apex, while on non-fruiting trees, they formed mainly from axillary buds, but in both cases, they tended to develop on younger parent shoots. The previous bearing condition of the tree was the main determinant for subsequent inflorescence development, which was independent of both shoot type and length. Thus, reproductive behavior strongly affected both the amount and type of new branching, but subsequent flowering level was more influenced by previous bearing than by the potential flowering sites on new shoots.     

Annual shoot-growth in Nothofagus antarctica (G. Forster) Oersted (Nothofagaceae) from northern Patagonia

The growth dynamics of annual shoots of Nothofagus antarctica shrubs from northern Patagonia was studied. The pattern of extension of the shoots derived from the three most distal buds of 100 parent shoots was registered. Weekly measurements and observations of extending shoots were carried out during the 1996–1997 growth season. The date of shoot extension initiation was less variable than the date of shoot extension cessation. Extension curves had an asymmetrically sigmoid outline. Most extension took place uninterruptedly between mid-September 1996 and the end of January 1997. Extension rate reached its highest value in the second half of the extension period. Leaf unfolding rate had a peak early in the extension period and fluctuated irregularly later on. The apical meristem of all shoots died by the end of shoot extension. In general terms, shoot length, diameter, leaf number and extension duration decreased from the first to the third position from the parent shoot’s apex, although exceptions to this pattern were found. Shoot extension rate was affected by maximum and minimum daily temperatures but not by the amount of precipitation during the extension period. The size of a bud was not related to the size of the shoot derived from it. The size of shoots in the third position from the apex was related to parent shoot size; this was not the case for more distal shoots. Received: 14 May 1999 / Accepted: 10 November 1999     

不同群落冠层环境下紫耳箭竹笋期生长发育研究

对金佛山国家级自然保护区内3个不同类型群落(落叶阔叶林、常绿落叶阔叶混交林、常绿阔叶林)下紫耳箭竹(Fargesia decurvata J. L. Lu)幼笋的生长发育进行研究。结果显示:(1)落叶阔叶林和常绿落叶阔叶混交林下的紫耳箭竹出笋期早而长,历时110 d,出笋量大,出笋率高;常绿阔叶林下的出笋期晚而短,历时88 d,出笋量少,出笋率低;常绿落叶阔叶混交林中出笋量最大。(2)出笋期分为3个阶段:初期、盛期和末期,各群落中的紫耳箭竹进入每个时期的时间有所差异,常绿落叶阔叶混交林中最早进入出笋盛期,落叶阔叶林次之,常绿阔叶林最迟。出笋盛期也是退笋的高峰期,退笋率的大小为:常绿落叶阔叶混交林落叶阔叶林常绿阔叶林。(3)同一群落冠层环境下,紫耳箭竹不同时期出土的幼笋地径无显著差异。在落叶阔叶林和常绿落叶阔叶混交林林冠环境中,各时期出土的幼笋地径间无显著差异,但均显著大于常绿阔叶林(P 0.05)。(4)紫耳箭竹幼笋出土后80 d左右完成高生长过程,且符合Logistic方程,呈"慢-快-慢"的生长趋势。高生长速率为:常绿落叶阔叶混交林落叶阔叶林常绿阔叶林,且差异显著(P 0.05)。(5)紫耳箭竹的克隆繁殖与分株密度间有密切关系。随着分株密度的增加,出笋数量增加,成竹数量降低。本研究表明不同群落冠层环境下紫耳箭竹生长发育存在显著差异,在常绿落叶阔叶混交林中发育最好,常绿阔叶林中发育最差,种群密度对竹类的更新发展起重要调节作用。     

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。     

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.     

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.     

EFFECTS OF AGE AND SIZE ON LIFE HISTORIES AND POPULATION GROWTH OF RHODODENDRON MAXIMUM SHOOTS

Age- and size-specific shoot life histories were studied with population censuses in June 1984 and June 1985 in an evergreen understory shrub. Rhododendron maximum. Most shoots (65%) survived without branching or flowering, and lesser numbers branched (2%), flowered (20%), or died (23%) during the year between censuses. The probabilities of surviving, branching, flowering or dying were both age- and size-dependent. Small, young shoots increased in leaf area. Flowering occurred most prominently in 3- to 6-year old shoots that had exceeded a leaf area of 200 cm², and the rate of flowering increased proportionately with size above this threshold. Branching normally occurred in the year following flowering. The age and size distributions of the population shifted significantly between years, indicating a nonequilibrium population. The survival schedule was Deevey Type I, indicating a high degree of “parental care” of young shoots. Age- and age + size-based demographic models predicted a rapid decline of the shoot population over a decade, while a size-based model predicted a much slower decline in shoot numbers. A sensitivity analysis of the models showed that overall shoot population growth was positively influenced by branching shoots and shoots that added leaf area, and negatively influenced by shoots that lost leaf area, died, or flowered. The role of shoot life histories in determining individual plant fitness and ecological dominance is discussed.     

Effects of shoot bending on ACC content,ethylene production,growth and flowering of bougainvillea

Several factors, such as environmental conditions, pruning, and plant growth regulators, affect the flowering of bougainvillea. However, information on the effect of shoot bending on growth and flowering of bougainvillea is scarce. In the natural environment, most of the bougainvillea flowering shoots are inclining whereas vertical shoots are not flowering shoots. Bougainvillea shoots are artificially grown vertically, horizontally and at an inclined orientation, to investigate the effect of these orientations on plant growth and the development of flower buds. The results of this indicate an effect of shoot bending on the growth rate of bougainvillea and the rate of flower bud formation. Additionally, our results suggest that vertical shoots have a higher growth rate, more prolific vegetation growth, and longer plastochrons (which are the intervals between the initiations of successive leaves). In contrast, horizontal and inclined shoots exhibited slower growth, a shorter time to reach flowering, and more flower buds. Inclined shoots had a higher endogenous ACC (1-aminocyclopropene-1-carboxylate) content and produced more ethylene than either horizontal or vertical shoots, indicating that more ACC in the inclined shoot is converted into ethylene, and the higher ethylene concentration in the inclined shoot causes it to mature earlier and flower sooner.     

千岛湖次生林优势种种群结构与分布格局

为了揭示亚热带地区常绿阔叶林次生演替机理,以砍伐后自然恢复40余年的浙江千岛湖姥山岛次生林为研究对象,分析了该群落优势种的种群结构、存活曲线和空间分布格局。结果表明:1)群落中重要值最大的是马尾松(Pinus massoniana)种群,达到50.47%,其后依次是石栎(Lithocarpusglabra)、苦槠(Castanopsis sclerophylla)和青冈(Cyclobalanopsis glauca)。2)马尾松的种群结构呈纺锤型,但1、2径级个体数极少,幼龄个体储备严重不足,已呈衰退趋势;石栎、苦槠、青冈的种群结构呈金字塔型。3)从静态生命表看,马尾松种群小径级的死亡率为负值,也说明马尾松种群幼苗严重缺乏,种群呈衰退趋势。4)马尾松种群的存活曲线为凸型,石栎、苦槠和青冈种群则近似直线型。5)4种优势种种群总体和各级个体的空间分布格局均呈聚集分布。总体来看,马尾松种群为稳定型趋向衰退型,而石栎、苦槠和青冈种群为稳定增长型,群落会继续向针阔混交林方向演替,进而演替成为常绿阔叶林。     

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.     

Correlative Inhibition in the Shoot of Agropyron repens ( L.) Beauv

Correlative inhibition was investigated in plants of Agropyronrepens at two temperatures. Reciprocal inhibition ocrurred betweenthe main shoot apex and the outgrowing axillary shoots, withthe balance of inhibition varying with temperature. Apical dominancewas stronger at 10 °C than at 20 °C , but even at 10°C release of apical dominance by decapitation had onlyminor effects on the timing of outgrowth, growth pattern andrate of dry weight aocumulation of the axillary shoots. Dominanceof the main shoot apex by the axillary shoots was stronger at20 °C than at 10 °C. Removal of axillary buds preventeddecline in size and activity of the main shoot apex ard resultedin increased rates of primordium initiation, leaf emergenceand dry weight accumulation in the main shoot. It is suggestedthat a system of reciprocal dominance provides a mechanism formaintaining the characteristic habit of the grass plant andlimits growth in height of vegetative shoots. Agropyron repens (L.) Beauv, couch grass, correlative inhibition, apical dominance, shoot, apex     

Spatial variation in polyphenolic content of Ascophyllum nodosum (Fucales,Phaeophyta)

Spatial variation in polyphenolic content in annual shoots of the brown alga Ascophyllum nodosum was quantified using a hierarchical sampling design. Three sampling levels, covering distances of 10⁰–10⁶ m, were used. Comparisons were made between two areas, Tjärnö on the Swedish west-coast and the Isle of Man in the Irish Sea, with very different types of environmental conditions. No significant differences in mean polyphenolic levels were found between the two study areas (6.6% of dry mass at Tjärnö and 9.2% at the Isle of Man), whereas significant and substantial differences were found among sites within areas (range 5.7%–11.4%) and among quadrats within sites (range 3.7%–13.1 %). The extensive variation at the smaller spatial scales points out the importance of using thorough sampling procedures at all levels in large-scale studies on algal polyphenolics, e.g. biogeographical comparisons, which have been neglected in several previous studies. Moreover, the results imply that experiments on causal factors of polyphenolic variation should be designed to explain the spatial scales on which the factors are impportant. This study also investigated the relationship between polyphenolic concentration and both plant size and mean area of annual shoots. The mean area was used as an estimate of the mean growth rate of the annual shoots within an individual. No significant relationships were found between shoot growth rate, or plant size, and polyphenolic levels in annual shoots at any of the three spatial scales that were investigated.     

太白山红桦种群结构与空间分析

 为了揭示太白山红桦(Betula albo-sinensis)种群的维持机制, 结合径级结构、静态生命表和存活曲线, 对太白山红桦种群结构进行了研究; 并应用单变量、双变量K函数对不同径级立木与残干的空间格局、空间关联性进行了多尺度分析。研究显示, 太白山红桦种群在幼苗幼树阶段死亡率较高, 进入中龄期后死亡率有所降低, 随着年龄增长, 死亡率逐渐增加, 直至年龄极限。其存活曲线基本接近Deevey I型(曲线凸型)。其个体死亡方式主要为掘根和折干(由大雪所致), 能够为种群更新提供潜在空间。除海拔2 250 m立木整体格局呈集群分布外, 该地区红桦立木与残干在不同尺度下的整体格局均为随机分布。不同发育阶段的个体均呈集群分布, 表现为斑块状同龄群。在红桦更新所需的最适林窗大小尺度上, 红桦立木与残干的空间关联性为显著正相关。以上结果表明: 太白山红桦种群具有同龄集群生长的特性, 种群由时空上不连续的局部斑块同龄个体组成, 其更新有赖于以掘根及折干方式死亡个体所形成的林窗斑块, 这种以局部林窗斑块更新维持种群整体稳定的特性, 可能是其长期稳定存在的重要机制。