Edge Effects and Seedling Bank Depletion: The Role Played by the Early Successional Palm Attalea oleifera (Arecaceae) in the Atlantic Forest

In this study, we examined the impacts of Attalea oleifera on the structure of seedling bank and discuss potential mechanisms of palm influence. Seed rain, seedling bank, and palm leaf fall were assessed beneath the canopy and in the vicinity of 16 adult palms across the edges of a large fragment (3500 ha) of the Atlantic forest. Moreover, we examined A. oleifera impacts on seed germination and seedling mortality by experimentally submitting seeds and seedlings to prolonged palm-leaf covering. As expected, seedling bank beneath the adults exhibited reduced abundance and species richness at local and habitat scale. Small to large seeds (3.1–30 mm) were underrepresented in the seed rain below adults palms, while experimental leaf covering drastically reduced both seed germination and seedling survivorship. A. oleifera leaf fall occurred over the whole year (3.6±2.7 leaves/individual/yr), which resulted in deep leaf litter mounds (10.7±9.2 cm). Finally, adult palm density (21.6±11.9 individuals/ha) correlated negatively with seedling density across Attalea clusters. Our results suggest that A. oleifera exerts negative effects on the seedling bank by reducing seedling abundance and richness as a consequence of two complementary mechanisms: impoverished and size-biased seed rain plus reduced seed germination and increased seedling mortality due to prolonged covering by fallen leaves.     

Responses of maternal siblings ofPinus densiflora to simulated add rain

Because most research on acid rain does not consider the genetic composition of seeds and the inherent variations in seed mass and germination date, the potentially diverse patterns of plant response are neglected within a population. Therefore, we examined the effects of both family identity and simulated acid rain onPinus densiflora seedlings over a 34-week period, beginning at germination. We were able to evaluate the importance of family and pH treatment independent of seed mass and germination date by considering the latter two variables as covariates. Seeds were collected from six randomly chosen trees that were assumed to represent different genotypes. After germination in a controlled environment, the seedlings were moved outdoors. Each was then subjected twice a week to 40 mL of simulated acid rain, at a pH of either 3.0 or 5.6. We examined variations in germination date, seedling height, needle numbers and lengths, and root and shoot masses. The maternal family effect was significant for all characters over the entire test period. Strong effects of pH treatment were detected only at the later growth stages. Artificial rain with a pH of 3.0 enhanced growth rates, particularly aboveground, compared with seedlings treated with pH 5.6 rain. However, regardless of pH, certain groups of maternal siblings consistently performed better in almost all parameters. The exception was for mean germination date, which differed among families only at a pH of 3.0. Therefore, although we demonstrated that acid rain affected both the date of germination as well as seedling growth rates inP. densiflora, the responses differed among maternal families, and were also associated with seed mass and germination date.     

Seed rain and soil seed bank of Picea asperata Mast. in subalpine coniferous forest of western Sichuan,China

Abstract

Dragon spruce (Picea asperata Mast.) is widely planted on clear-cuts from natural subalpine coniferous forests in western Sichuan. To assess the natural regeneration potential of this species in spruce plantations of different ages, field studies on the seed rain, seed bank, and seedling recruitment were conducted in 20-year, 30-year, and 60-year plantations, and in a retained natural forest ca. 150 years old for comparison. Moreover, a series of temperature and light regimes were also designed in March 2003 to test germination/dormancy responses of the P. asperata seeds to different conditions. In the plantations considered, both the densities of seed rain and soil seed bank increased with increasing stand age, whereas they were both low beneath the natural forest partially due to low adult density of P. asperata adult trees. P. asperata has a transient seed bank, and ca. 60% of seeds were found in the litter layer. Seed decay and seed predation were the two most important factors affecting soil seed bank dynamics, which caused a number of seed losses from the soil. Only a small fraction germinated and produced seedlings, and the indices of the losses from the seed bank via germination in the 30-year plantation, 60-year plantation and natural forest were 4.75, 5.10, and 2.80%, respectively. High seedling mortality was observed after seedling emergence, and most of the germinated seedlings died out within one growing season. The P. asperata seeds showed a high percent germination and no significant differences under most light and temperature regimes except for full sunlight or temperature below 5°C. In conclusion, despite a substantial number of seed produced, the high depletion of soil seed, the low seedling output, and high seedling mortality may obscure the natural regeneration potential of this tree species.     

Seedling emergence patterns and dormancy/germination physiology of <Emphasis Type="Italic">Primula modesta</Emphasis> in a subalpine region

On the basis of the germination/dormancy responses of seeds to temperature and light, and local seed rain, we attempted to interpret the seedling emergence patterns of Primula modesta Bisset et Moore (Primulaceae) in two different types of habitats in a subalpine zone of Mt. Asama: an oligotrophic flat moor and a grassland with relatively dense herbaceous vegetation cover. The seasonal pattern of seedling emergence was well explained by the dormancy/germination physiology revealed in laboratory germination tests. The seeds were demonstrated to have a strict light requirement even after experiencing moist chilling, which might facilitate the incorporation of the seeds into the soil seed bank. Despite sufficient seed production, the seedlings emerging were far less at the grassy site than the moor site, but the number of seedlings was significantly dependent on the seed rain within previous season and on the litter cover of the microsite in both sites. Therefore, the spatiotemporal patterns of seedling emergence in the habitats could be well explained by the spatiotemporal patterns of seed rain and safe-sites for germination.     

Regenerative traits of tree species in a deciduous forest of northeastern North America

I studied the seed rain, seedling bank, newly emerged seedlings, and seed bank of tree species in a deciduous forest of northeastern North America (Québec, Canada). The main objective was to determine to what extent the tree species present on the site differed in some of their regenerative traits. All the species present on the quadrats as mature trees contributed to the annual seed rain, although seed rain was strongly dominated by yellow birch and sugar maple. The seedling bank was mostly composed of red maple and sugar maple, but inputs from the 1988 cohort were mostly red maple and yellow birch seedlings. Seedling mortality was high in general (from 60% to 100%), but sugar maple had the lowest mortality of the tree species present. Two species maintained a seed bank on the study site: yellow birch and american basswood. Some species not present on the quadrats, but having populations nearby, disseminated seeds onto the quadrats (i.e. white ash and grey birch). Other species, not present on the quadrats as mature individuals, had seeds in the seed bank although they did not participate in the seed rain, did not maintain a seedling bank, and did not establish seedlings in 1988 (i.e. pin- cherry and Rubus spp.). Red maple, sugar maple, and yellow birch showed contagion in the dispersion pattern of their seeds in the seed rain (and the seed bank for yellow birch), and of their seedlings in the seedling bank and the 1988 cohort. Seedlings of american beech and american basswood of the seedling bank and the 1988 cohort were randomly dispersed over the quadrats, although their seeds in the seed rain had a contagious dispersion pattern. It has often been proposed in the literature that regenerative traits come only in “compatible assemblages” and that, consequently, species may be grouped according to their strategy of regeneration: e.g. species that maintain a seed bank vs those with a seedling bank; species with either a seed or a seedling bank vs those with the ability to sprout. In the community studied, it appears that regenerative traits do not come in neatly packaged exclusive assemblages, although the species do seem to possess different combinations of regenerative traits. However, yearly variations in seed production, seed germination, and seedling survival may render difficult the identification of strategies of regeneration.     

The dynamics of three shrub species in a fire-prone temperate savanna: the interplay between the seed bank, seed rain and fire regime

A model was developed to assess how the seed rain and fire regime affect seed bank dynamics and seedling establishment of three native shrub species (Acanthostyles buniifolius, Baccharis pingraea and Baccharis dracunculifolia) with different regeneration strategies, in temperate South American savanna. Seed bank and seed rain were quantified for each species under different fire regimes, and their relative roles in regeneration were evaluated. All species had short-term persistent seed banks and high annual variability in seed production. A high proportion of seeds deposited in the seed rain produced seedlings after fire; few entered the soil seed bank. Fire killed a high proportion of the seeds in the soil seed bank. Seedlings derived from the seed rain had a higher probability of surviving for 2 years than seedlings emerging from the soil seed bank. In the absence of fire, establishment depended on germination both from the seed rain and the soil seed bank, whereas with annual fire, establishment was primarily dependent on germination of seeds arriving in the annual seed rain, regardless of species’ regeneration strategies. These results help to explain changes in the vegetation of South American temperate savannas as a result of changes in fire regime and grazing management during the last 50 years. By revealing the crucial roles of the soil seed bank and seed rain in regeneration, this study provides vital information for the development of appropriate management practices to control populations of shrub species with different regeneration strategies in South American temperate savannas.     

模拟秋季酸雨对三角枫叶片光合生理特性的影响

以三角枫幼苗为材料,采用盆栽方法,研究了pH 5.6、pH 4.0、pH 3.0和pH 2.0酸度模拟酸雨胁迫对三角枫叶片光合生理特性的影响,以探讨酸雨胁迫下三角枫的光合生理响应机制。结果显示:(1)随着酸雨浓度的增加和胁迫时间的延长,三角枫叶片相对叶绿素含量下降幅度逐步增大;叶片丙二醛含量呈逐渐上升的趋势,且各处理均显著高于对照;叶片质膜透性和脯氨酸含量均呈先升高后降低的趋势,且pH 2.0处理的叶片质膜透性在试验20d时迅速增大,升高幅度最大(146.3%)。(2)叶片净光合速率、蒸腾速率、水分利用效率、表观光能利用效率以及表观CO2利用效率在胁迫下也均呈显著下降趋势,且以pH 3.0和pH 2.0处理降幅最大。研究表明,pH 4.0的模拟酸雨对三角枫叶片的光合生理指标无显著影响,而pH≤3.0的强酸雨胁迫使三角枫叶片叶绿素含量显著下降、膜保护系统受损、光合作用效率显著下降;三角枫能适应弱酸雨(pH≥4.0)环境,可作为酸雨地区的园林绿化树种。     

Regeneration filters in post-agricultural fields of Puerto Rico and Ecuador

To better understand how rainforest regenerates after agriculture I sampled the seed rain and seed bank, and set out seeds and seedlings, on microsites defined by distance to the forest in fields both in Puerto Rico and Ecuador. I found that (1) total seeds, species richness and life-form richness were twice as great in the Ecuador seed rain compared to Puerto Rico but Puerto Rico seeds were more evenly distributed among species and (2) total seedlings from the seed bank were similar between Puerto Rico and Ecuador, (3) the majority of seeds were lost to predation among all species and study sites, (4) seed disease was absent in P. aduncum and Miconia prasina, and no seeds germinated for Gonzalagunia spicata and P. riparia, (5) in Ecuador pathogenic disease claimed more seeds than germinated for all species, and Solanum ovalifolium was the only species that had seeds germinate but did not lose seeds to disease, (6) also in Ecuador, insect predation was significantly lower in the forest border for P. aduncum, and seed disease was significantly greater at the 10 m micro site for S. ovalifolium, (7) distance has a significant effect on seedling height and basal diameter, (8) losses of leaf area due to herbivory and pathogens were always low and (9) biomass and leaf specific mass were significantly reduced in the border and forest microsites. I conclude that Ecuador fields had more seeds, species, and life-forms than Puerto Rico fields, predation was the most severe post-dispersal seed filter in all fields, seeds that survived predation on Puerto Rico were lost either to disease or germination but to both mechanisms in Ecuador, all three seed mechanisms in Ecuador fields showed distance effects of seedling growth but not survivorship.     

模拟酸雨对西洋杜鹃生理生态特性的影响

采用盆栽方法,研究了不同pH值条件下的模拟酸雨对西洋杜鹃生长及其叶片主要生理生化特征的影响。结果表明:在酸雨胁迫下,随着酸雨胁迫的增强,西洋杜鹃叶片受害程度逐渐加重;叶片丙二醛含量逐渐升高;可溶性蛋白含量先升后降;过氧化氢酶(CAT)、过氧化物酶(POD)和超氧化物歧化酶(SOD)活性均呈先升后降的单峰曲线变化,其中pH值为4.3的处理下西洋杜鹃叶片CAT和SOD活性最高,pH值为3.0的处理下其POD活性最高;其叶绿素含量逐渐下降;叶片净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)及水分利用率(WUE)均呈先升后降的趋势,而胞间CO2浓度(Ci)则持续下降,pH值为4.3的处理下其Pn、Tr、Gs和WUE均达到最高。研究表明,大致可以认为pH值≤3.0是酸雨对西洋杜鹃造成隐形伤害的阀值,而酸雨灾害严重地区的降水pH值为2.0—4.0左右,说明西洋杜鹃可以在酸雨灾害较重的地区生长,可作为酸雨灾害严重地区园林绿化及植被构建的物种之一。     

毛红椿天然林种子雨、种子库与天然更新

2008-2011年,调查江西九连山国家级自然保护区毛红椿天然林的种子雨、种子库及林下幼苗数量.结果表明:在毛红椿天然林,种子雨散布时间为10月下旬至12月下旬.2010年不同样地的种子雨强度为虾公塘气象观测站(320.3±23.5粒·m-2)＞虾公塘保护站(284.7±24.2粒·m-2)＞大丘田保护站(251.6±24.7粒·m-2),分别以222.0、34.3和22.6粒·m-2完好种子供土壤萌发更新;毛红椿种子储量取决于结实量、鸟类取食和种子活力等因素,鸟类取食是其种子储量大幅下降的首要因素;由于种子不耐储藏以及大量腐烂,种子有效贮藏期不足1个月.12月天然林种子库平均萌发数≤2株·m-2,次年1月土壤种子库种子量最少,为6.7～11.8粒·m-2,平均仅萌发0.4～0.6株·m-2,与林下实生幼苗分布极少相吻合.毛红椿种子雨储备、种子库种子活力保存及幼苗建成等因素影响其天然更新.     

文峪河上游华北落叶松林的种子雨、种子库与幼苗更新

华北落叶松林下更新不良,为探究其制约因素,开展了山西省文峪河上游5个华北落叶松林分的种子雨、土壤种子库和幼苗更新的研究。结果表明:(1)华北落叶松种子主要集中于9—10月散落。2011年为华北落叶松种子丰年:种子产量高,种子雨密度达(961.93±377.40)粒/m2;种子质量高,完整种子占(89.31±16.13)%。2012年为种子平年,种子产量低,种子雨密度为(252.73±115.12)粒/m2。华北落叶松种子雨主要源于毗邻树木,华北落叶松纯林和落叶松云杉林的种子雨密度显著高于其他3个针阔混交林。(2)土壤种子库主要由上年种子雨组成,2012年4月的土壤种子库密度为(695.18±297.23)粒/m2,完整种子占(59.73±9.56)%。种子自然萌发前,约(78.98±24.76)粒/m2具发芽力,基本可满足更新需要。但种子活力保持期少于2 a,只能形成短期持久土壤种子库。(3)华北落叶松更新不良,种子年后仍难以实现幼苗建成,当年生幼苗的出现频度平均为1.6%,且林下难以存活。幼苗发生与种子储量关联性不强,种源条件不是制约华北落叶松更新的主要因素。     

不同年龄香果树种子雨和种子库及其更新特征

对宝天曼自然保护区香果树种子雨、土壤种子库进行了观测,并进行不同微生境播种实验,研究其种子萌发及幼苗生长动态。结果表明:(1)不同龄级香果树种子雨持续时间及其高峰期有所不同,种子雨强度存在极显著差异。(2)20~50龄级香果树的种子饱满率、土壤种子库中种子密度均显著低于其他龄级,但其不同类型种子的水平和垂直分布规律一致;大约80%的香果树种子在其萌发前消失,剩余种子中以霉烂种子密度最高(9.81粒·m-2),饱满种子密度最低(1.94粒·m-2)。(3)野外育苗实验表明,不同龄级香果树所产饱满种子的萌发率及幼苗存活率差异不显著,其种子萌发率约为16.93%,但仅有3.86%的幼苗寿命超过5个月;不同微生境对香果树幼苗存活率产生显著影响,林窗是其最适宜微生境。研究认为,人工采集香果树种子,于次年4月份在原生境下播种;清理母树冠下和冠缘的地被物,增加林缘空地地被物覆盖度,以减少香果树种子的损失,改善香果树种子萌发和幼苗生长的光照条件,以利于幼苗根系的生长,促进香果树的自然更新。     

海南霸王岭陆均松种子雨和土壤种子库特征

为了解陆均松(Dacrydium pierrei)的自然更新能力,对海南霸王岭陆均松进行为期3年的种子雨、种子库及幼苗幼树观测,并分析其与环境因子的关系。结果表明,陆均松种子雨和种子库有效种子数量太少并存在季节性差异;其结实存在大小年现象;种子雨的扩散密度随着距母树距离的增加而逐渐减少;土壤种子库种子数量在枯枝落叶层所占比例最大,但有活力种子数量极少。幼苗幼树适宜生存的海拔范围为800~1 400 m,经度、纬度、坡度与幼苗幼树分布有明显相关性。因此,选择适宜季节采摘种子,提高种源数量和质量,清除林下枯枝落叶,增加种子萌发能力等以减少陆均松更新的限制条件的人工抚育措施应适当考虑。     

Disperser- vs. Establishment-Limited Distribution of a Riparian Fig Tree (Ficus insipida) in a Costa Rican Tropical Rain Forest1

Studies were conducted at the La Selva Biological Station in Costa Rica and in a greenhouse in California to determine the factors accounting for the nonrandom distribution of the riparian fig tree Ficus insipida Willd. along streams in the La Selva Biological Reserve and adjacent deforested lands. We also evaluated the potential seed dispersers of this tree relative to the role of the fruit‐eating fish Brycon guatemalensis that previously was proposed to be an important disperser of F. insipida seeds in this system. At La Selva, we recorded the fig‐foraging activities of vertebrates at fruiting F. insipida trees, surveyed for the presence or absence of F. insipida along streams of different sizes, and determined the fate of fig seedlings transplanted in different riparian habitats. In the greenhouse, we measured seed germination and seedling survival and growth under different light and soil pH conditions mimicking natural conditions. The findings provided evidence that (1) the tree occurs along the larger streams running through forest habitat and only along smaller streams with relatively high light availability; (2) bats (Artibeus spp.) and fish are the major dispersers of F. insipida seeds; (3) the seedlings are subject to mortality not only from low light conditions but also from treefalls, frequent flooding, and bank erosion; and (4) high light levels and near neutral soil pH result in relatively better seed germination, faster growth, and higher survival rates of seedlings. Overall, our results suggest that this fig tree is dispersed mainly by bats and fish and is more establishment‐limited than disperser‐limited in its local distribution in the La Selva rain forest habitat.     

Ten-year seed bank and vegetation dynamics of a tidal freshwater marsh

A decade-long examination was made of recruitment and establishment in a tidal freshwater high marsh along the Delaware River. Over the 10 yr of the study, seed bank, field seedlings, and vegetation showed variable patterns and significant year-to-year fluctuations. Patterns of each species were unique, perhaps the result of specific germination and/or establishment requirements and seedling morphology. For a given species there was little correlation among seed bank, seedling, and vegetation patterns, and germination success did not guarantee establishment. Species diversity showed significant year-to-year fluctuation, but there was no trend, and perennials did not change in importance during the 10-yr period. Because four annual species (Bidens laevis, Cuscuta gronovii, Impatiens capensis, and Polygonum arifolium) composed over 90% of the seed bank and field seedlings, and 58-89% ( = 70 ± 4) of the cover, community dynamics were dependent on seedling recruitment. For a given species life history stages (seed bank, field seedlings, and vegetation) were not predictable over the temporal scale of a decade. It is predicted, however, that if hydrology remains unchanged, the same suite of species will persist. The importance of the parasite Cuscuta gronovii is noteworthy.     

六盘山区辽东栎的实生苗更新及其影响因子

研究了六盘山区辽东栎(Quercus liaotungensis)灌丛的种子产量、土壤种子库组成、湿沙和风干贮藏对种子寿命的影响以及动物取食子叶对种子萌发和幼苗建立的影响。结果表明, 辽东栎灌丛种子的完好率为27.51%, 被动物取食或搬运种子的比例(41.51%)显著高于其他类型种子(p < 0.01); 辽东栎次生林土壤种子库中萌发和虫蛀种子分别占35.16%和38.29%, 完好种子仅占13.65%, 捕食动物主要通过贮藏或搬运而影响土壤种子库中的种子密度。湿沙贮藏60天的辽东栎种子自动萌发率高达96.67%, 短期贮藏可加快种子的萌发进程, 提高萌发率和萌发指数, 但随着贮藏时间的延长, 种子萌发进程延迟, 萌发率、萌发指数和活力指数均不同程度地降低。排除动物取食处理的幼苗在林窗和林下生境的存活率分别为80%和83%, 而不排除动物取食幼苗在2种生境中分别仅有25%和31%能够存活, 表明子叶在幼苗建立中具有重要作用。林窗中幼苗子叶的动物取食率(85.00%)高于林下(71.00%), 子叶留存的幼苗在林窗中的存活率(6.00%)低于林下(15.50%), 而子叶被取食幼苗的存活率在两种环境中基本相等(分别为18.50%和18.00%)。     

盐胁迫下赤霉素对黄瓜种子萌发及幼苗生长的影响

研究外源GA₃对盐胁迫下黄瓜种子萌发和幼苗生长的影响。结果表明,添加不同质量浓度GA₃的各处理,其发芽率、发芽势和发芽指数均显著高于NaCl胁迫处理,其中以100 mg/L GA₃处理的发芽势、发芽率和发芽指数最高,幼苗的叶面积、根长、根冠比也最大,同时叶片中叶绿素含量最高,幼苗叶片的光合速率(P_n)、气孔导度(G_s)、胞间CO₂摩尔分数(C_i)及蒸腾速率(T_r)等均达到最大;而当赤霉素的质量浓度为50 mg/L时,叶片中的POD活性为2 005 U/(g·min）,达最大值。     

金丝李果实、种子形态分化及外源物质对种子萌发和幼苗生长的影响

通过采集3个天然种群的金丝李(Garcinia paucinervis)果实,观察果实和种子的形态性状,分析其种群内和种群间的形态分化,观察不同种群种子萌发和幼苗生长的规律;采用四种植物生长调节剂和两种化学药剂,研究浸种对金丝李种子萌发和幼苗生长的影响。结果表明:金丝李种子长2.48~3.08 cm、宽1.49~1.67cm、百粒重392.50~438.18 g;3个种群果实和种子已产生明显的形态分化,胡润种群多数表型性状值在3个种群中最小,安宁种群的果实和种子偏圆,弄岗种群偏窄;在种群内部,果实和种子重量的分化较显著,整体形态较稳定;在萌发过程中,种子留存,不定根逐渐取代胚根成为主根;3个种群的萌发过程均缓慢,中间阶段几乎停滞,出苗极不整齐,萌发率表现为安宁种群(78.33%)弄岗种群(55%)胡润种群(48.33%),种子存在休眠;从种子萌发来看,以80 mg·L~(-1)的6-BA浸种24 h处理效果最好,萌发时滞最短、萌发率最高、平均萌发时间最短;结合幼苗生长情况来看,10~50 mg·L~(-1)的6-BA和500 mg·L~(-1)的GA_3处理均适宜,以50 mg·L~(-1)的6-BA效果最佳。低浓度的KNO_3和NH_4NO_3能提高萌发率,但未显著加快萌发进程,IAA和NAA对种子萌发和幼苗生长有抑制作用。该研究结果为金丝李的遗传多样性提供了形态学资料,为更好地保护、开发和利用这一珍贵树种提供了科学依据。     

The effect of water-level fluctuations on seedling recruitment in an aquatic macrophyte Nymphoides indica (L.) Kuntze (Menyanthaceae)

We examined seed bank characteristics and seed germination and the survival of seedlings to understand the significance of water-level fluctuations on sexual regeneration in Nymphoides indica (L.) Kuntze. The seed bank existed throughout the pond. The longevity of the seed bank was more than 3 years. Seed germination started on the exposed mudflat as soon as water drawdown occurred. At the juvenile stage, exposed conditions were required for seedling growth and the young seedlings died under submergence. It was suggested that seedling establishment was successful after occasional water drawdown or along the water margin according to the period and pattern of water-level fluctuation. This study indicated the significance of the exposed mudflat at the water edge for seed germination and the growth of juvenile plants to the success of sexual regeneration in N. indica.     

Regeneration by seeds in alpine meadow and heath vegetation in sub‐arctic Finland

Abstract. This paper compares the regeneration by seeds of heath and meadow and studies relationships between the floristic composition of phases in the regeneration pathway. Seed densities in the seed rain and seed bank as well as the densities of emerged seedlings in gaps and in closed vegetation were greater in the meadow than in the heath. In the heath, environmental constraints hindered seedling emergence almost completely so seeds accumulated in the seed bank. In the meadow, the decrease in the seed bank was due to high seedling emergence. Within both plant communities, seedling emergence in gaps and in closed vegetation was comparable. In the meadow, the seed rain and seedling emergence in gaps, as well as the seed bank and seedling emergence in gaps were positively correlated. Differences in seed and adult plant sizes were reasons for the low correlation between the standing vegetation and the other phases. In DCA ordination the first axis separated the phase of seedling emergence in closed vegetation and seed bank. The second axis separated the standing vegetation from the other phases. The structure of the seed rain was more heterogeneous than that of other phases. In the heath, the standing vegetation and the seed rain were positively correlated. The ordination of these phases reflected the patchiness of standing vegetation and the ability of the diaspores of Betula nana to disperse over long distances.