Density-dependent regulation of ramet recruitment by the red:far-red ratio of solar radiation: a field evaluation with the bunchgrass Schizachyrium scoparium

Depressions in the red to far-red ratio (R:FR) of solar radiation arising from the selective absorption of R (600–700 nm) and scattering of FR (700–800 nm) by chlorophyll within plant canopies may function as an environmental signal directly regulating axillary bud growth and subsequent ramet recruitment in clonal plants. We tested this hypothesis in the field within a single cohort of parental ramets in established clones of the perennial bunchgrass, Schizachyrium scoparium. The R:FR was modified near leaf sheaths and axillary buds at the bases of individual ramets throughout the photoperiod without increasing photosynthetic photon flux density (PPFD) by either (1) supplementing R beneath canopies to raise the naturally low R:FR or (2) supplementing FR beneath partially defoliated canopies to suppress the natural R:FR increase following defoliation. Treatment responses were assessed by simultaneously monitoring ramet recruitment, PPFD and the R:FR beneath individual clone canopies at biweekly intervals over a 12-week period. Neither supplemental R nor FR influenced the rate or magnitude of ramet recruitment despite the occurrence of ramet recruitment in all experimental clones. In contrast, defoliation with or without supplemental FR beneath clone canopies reduced ramet recruitment 88% by the end of the experiment. The hypothesis stating that the R:FR signal directly regulates ramet recruitment is further weakened by evidence demonstrating that (1) the low R:FR-induced suppression of ramet recruitment is only one component of several architectural modifications exhibited by ramets in response to the R:FR signal (2) immature leaf blades, rather than leaf sheaths or buds, function as sites of R:FR perception on individual ramets, and (3) increases in the R:FR at clone bases following partial canopy removal are relatively transient and do not override the associated constraints on ramet recruitment resulting from defoliation. A depressed R:FR is probably of greater ecological significance as a signal of competition for light in vegetation canopies than as a density-dependent signal which directly regulates bud growth and ramet recruitment.     

Light limitation creates patchy distribution of an invasive grass in eastern deciduous forests

Species interactions and their indirect effects on the availability and distribution of resources have been considered strong determinants of community structure in many different ecological systems. In deciduous forests, the presence of overstory trees and shrubs creates a shifting mosaic of resources for understory plants, with implications for their distribution and abundance. Determination of the ultimate resource constraints on understory vegetation may aid management of these systems that have become increasingly susceptible to invasions by non-native plants. Microstegium vimineum (Japanese grass) is an invasive annual grass that has spread rapidly throughout the understory of forests across the eastern United States since it was first observed in Tennessee in 1919. M. vimineum occurs as extensive, dense patches in the understory of eastern deciduous forests, yet these patches often exhibit sharp boundaries and distinct gaps in cover. One example of this distributional pattern was observed relative to the native midstory tree Asimina triloba (pawpaw), whereby dense M. vimineum cover stopped abruptly at the drip line of the A. triloba patch and was absent beneath the A. triloba canopy. We conducted field and greenhouse experiments to test several hypotheses regarding the causes of this observed pattern of M. vimineum distribution, including allelopathy, seed dispersal, light limitations, and soil moisture, texture, and nutrient content. We concluded that light reduction by the A. triloba canopy was the environmental constraint that prevented establishment of M. vimineum beneath this tree. Whereas overstory tree canopy apparently facilitates the establishment of this shade-tolerant grass, the interaction of overstory canopy with midstory canopy interferes with M. vimineum by reducing the availability of sunflecks at the ground layer. It is likely that other midstory species influence the distribution and abundance of other herb-layer species, with implications for management of understory invasive plant species.     

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]     

Genet structure and determinants of clonal structure in a temperate deciduous woodland herb, Uvularia perfoliata

1 We used isozyme variation to examine the genet structure of Uvularia perfoliata patches in gap and closed canopy habitats in a temperate deciduous forest in Maryland, USA.
2 A large patch in a gap habitat was composed of a small number of widely spread genets with many ramets, and a large number of genets with more restricted distribution and few ramets. Genets with many ramets were patchily distributed at a metre scale. Analysis of genet structure on a scale of square centimetres, however, revealed that the genets were highly intermingled with no clear boundaries between them. The presence at both scales of sampling of many genets with unique multilocus genotypes indicated continuing genet recruitment within the population.
3 In the closed canopy habitat, the patches examined were each composed of a single unique multilocus genotype, suggesting that each had developed by asexual propagation following the establishment of a single genet.
4 The clonal structure of U. perfoliata patches in both gap and closed canopy habitats therefore appears to depend on recruitment patterns of genets. Populations in closed canopy habitats are characterized by a 'waiting' strategy, in which asexual ramet production maintains populations until genet recruitment by seed production can occur under the more optimal conditions associated with canopy gaps. Extended sampling suggests that the genetic diversity of U. perfoliata populations is primarily controlled by the disturbance regime of the forest canopy.     

The regeneration niche of the grey mangrove (Avicennia marina): effects of salinity,light and sediment factors on establishment,growth and survival in the field

Field observations of seedlings and saplings of Avicennia marina showed patterns that correlated with salinity, light and sediment. Models that account for these observations were subsequently tested in a series of field experiments. Establishment varied within an estuary under controlled conditions but was not related to salinity or sediment type. Seedling survival was uniform over 3 years regardless of position in estuary and sediment type. Seedling densities and survival under canopies or in canopy gaps were not significantly different. However, seedling growth and density of saplings were greater in canopy gaps. Experiments involving manipulations of canopies showed no differences in seedling survival under canopies or in light gaps, but addition of slow-release fertilizer enhanced growth and survival in canopy gaps and under canopies. Long-term comparison of areas denuded of a canopy and with sediment disturbance showed enhanced establishment and survival when compared with areas with canopy gaps but with undisturbed sediments. Overall there appears to be no restriction to establishment of propagules within mangrove stands other than the supply of propagules and tidal or wave action. In contrast, recruitment to the sapling stage appears to be restricted by light and sediment resources. We suggest that propagules need to establish in a regeneration niche for seedling recruitment to the sapling stage. This differs from the view that seedlings in the under-storey are analogous to a seed pool in the soil.     

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 condi-tions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to dif-ferent 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 clus-ter 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. More-over, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heteerogeneous light environoments.     

THE RELATIONSHIPS BETWEEN SEASONAL VARIATION IN LIGHT INTENSITY,RAMET SIZE,AND SEXUAL REPRODUCTION IN NATURAL AND EXPERIMENTAL POPULATIONS OF ASTER ACUMINATUS (COMPOSITAE)

We conducted experiments to evaluate the effects of different patterns of seasonal light availability on ramet size and sexual reproduction in the understory herb, Aster acuminatus. In April 1982, 200 potted rhizomes were divided equally between a high light and a shaded site. Twice during the growing season subsets of pots were reciprocally transferred between the sites, resulting in eight different light treatments. There was significant variation among treatments in mean ramet weight (0.41–1.21 g), height (15.0–23.4 cm), number of flowers (0.4–7.9) and percentage of ramets flowering (14–95). Periods of high light increased ramet height, weight, rhizome production and flowering, but high light earlier in the season increased ramet height more than did high light later in the season, while high light later had pronounced effects on the other three traits. Measurements taken in nine natural patches experiencing different patterns of seasonal light availability are consistent with the experimental results. The results provide new insights into the general phenomenon of size-dependent reproduction in herbs and illustrate how the phenology of biomass allocation can affect growth and reproduction.     

Influence of elevated CO2 on canopy development and red:far-red ratios in two-storied stands ofRicinus communis

Vertical structure of plant stands and canopies may change under conditions of elevated CO₂ due to differential responses of overstory and understory plants or plant parts. In the long term, seedling recruitment, competition, and thus population or community structure may be affected. Aside from the possible differential direct effects of elevated CO₂ on photosynthesis and growth, both the quantity and quality of the light below the overstory canopy could be indirectly affected by CO₂-induced changes in overstory leaf area index (LAI) and/or changes in overstory leaf quality. In order to explore such possible interactions, we compared canopy leaf area development, canopy light extinction and the quality of light beneath overstory leaves of two-storied monospecific stands ofRicinus communis exposed to ambient (340 μl l⁻¹) and elevated (610 μl l⁻¹) CO₂. Plants in each stand were grown in a common soil as closed “artificial ecosystems” with a ground area of 6.7 m². LAI of overstory plants in all ecosystems more than doubled during the experiment but was not different between CO₂ treatments at the end. As a consequence, extinction of photosynthetically active radiation (PAR) was also not altered. However, under elevated CO₂ the red to far-red ratio (R:FR) measured beneath overstory leaves was 10% lower than in ecosystems treated with ambient CO₂. This reduction was associated with increased thickness of palisade layers of overstory leaves and appears to be a plausible explanation for the specific enhancement of stem elongation of understory plants (without a corresponding biomass response) under elevated CO₂. CO₂ enrichment led to increased biomass of overstory plants (mainly stem biomass) but had no effect on understory biomass. The results of this study raise the possibility of an important indirect effect of elevated CO₂ at the stand-level. We suggest that, under elevated CO₂, reductions in the R:FR ratio beneath overstory canopies may affect understory plant development independently of the effects of PAR extinction.     

Light-induced variation in the growth and dynamics of transplanted ramets of the understory herb,Aster acuminatus

Summary Ramets of the understory herb, Aster acuminatus, were transplanted from two source populations into eight understory garden sites that varied in light and soil moisture levels. Ramet growth, clonal growth, flowering and survivorship were monitored for three growing seasons. Large differences among gardens in ramet growth, clonal growth and flowering developed in the first growing season and increased in the next two years. This variation was positively correlated with garden light level but not at all with soil moisture. Mortality rates were low in all gardens and showed that genets from any particular source could survive over a broad range of environmental conditions. There was no conclusive evidence for any source population differences in the capacity to survive or grow in different environments. The rapid, light-induced responses of transplanted ramets resulted in garden populations very similar in appearance to natural populations experiencing similar light regimes. These results combined with those from other studies of A. acuminatus provide strong evidence for the importance of light in explaining population patterns and dynamics in this species.     

亚高山暗针叶林不同林冠环境下华西箭竹的克隆可塑性

以亚高山暗针叶林3种林冠环境中以及暗针叶林林缘的华西箭竹（Fargesia nitida）为对象,对其无性系数量特征、无性系根茎特征、分株生物量以及分株形态特征进行了对比研究。结果表明：（1）林冠环境的差异导致了不同种群的基株密度和每基株分株数的显著差异,但林冠环境差异不影响分株密度。林冠郁闭度愈大,每基株分株数愈少,分株分布愈均匀。（2）不同林冠环境间。分株生物量、分株构件生物量和分株构件的生物量分配百分率均有显著差异。开敞的林冠环境有利于华西箭竹的生长和生物量积累。（3）随着林冠郁闭度的增加,华西箭竹通过增大分枝角度、叶生物量分配百分率、比叶面积和叶面积率以提高光能利用效率,有效适应弱光环境。（4）隔离者长度、隔离者直径和分枝强度在林缘和林窗环境中要显著大于林内环境;同级隔离者分枝角度随林冠郁闭度的增加而最大,其值在林下显著大于林窗和林缘,而异级隔离者分枝角度的变化则正好相反。研究表明,华西箭竹种群在不同的林冠环境中发生了明显的可塑性变化,这些可塑性变化是种群对林冠郁闭度差异的适应性反应的结果,有利于增强种群对环境中有限光资源的利用。     

Species-specific sprouting pattern in two dioecious Lindera shrubs: The role of physiological integration

Sprouting is recognized as an important genet persistence strategy for clonal woody plants, but the role of sprouting may differ between species and between sexes, depending on physiological integration. We tested the effect of physiological integration on the mortality, recruitment and growth of the sprouting male and female ramets of two closely related dioecious shrubs of Lindera, in a field experiment using girdling manipulation. Although between-sex differences observed were obscure, we found between-species differences in the sprouting patterns. The rates of ramet mortality and recruitment were significantly lower for L. praecox than L. triloba. In L. praecox genets, the ramet production was low, and the main ramets might actively translocate assimilates towards the small sprouted ramets, which then facilitates high ramet growth and survival (sprout-nursing strategy). Meanwhile, in L. triloba genets, although many ramets were recruited, assimilate translocation from the main ramets to the sprouted ramets might be less abundant, which causes high ramet mortality (sprout-turnover strategy). For a more general knowledge of the various sprouting strategies in clonal plants, our study demonstrated that inter-specific comparisons using girdling experiments at the whole-plant level could reveal the role of physiological integration on the link between the sprouting pattern and above-ground structures of clonal plants.     

Internal heterogeneity of ramet and flower densities of <Emphasis Type="Italic">Typha angustifolia</Emphasis> near the boundary of the stand

Fifteen relatively narrow stands of Typha angustifolia, including both stands bounded to open water and ridged boundary, were investigated in terms of the distributions of ramet density as well as the fraction of flowering ramets. The ramet density, fraction of flowering ramet and morphological characteristics of ramets were measured continuously from one side of the stand to the other, together with depth and the nutrient concentration of pore water. The rhizome biomass was also measured at the periphery and at the center of the stand. The results indicated that the shoot density declined gradually from the edge to the center, however, the fraction of the flowering ramets were significantly higher at the edge while it rapidly declined away from the edge. Statistical analyses indicate that the ramet density depends on either depth or nutrient level of the pore water; however, the distance from the edge was the most probable factor to reduce the fraction of the flowering ramets. The rhizome biomass was mostly larger at the periphery than at the center, likely because the accumulation of the horizontal rhizomes as extension was blocked by the boundary. High flowering fraction near the periphery was probably induced by the hormonal change caused by the blockage. However, a large fraction of ramets flowered simultaneously with large rhizome biomass at the periphery, likely because of the preferable light climates compared with inside the stand.     

冠层结构和光环境的时空变化对紫耳箭竹种群特征的影响

选择金佛山国家自然保护区内落叶阔叶林、常绿落叶阔叶混交林、常绿阔叶林3种典型群落类型,研究冠层结构和光环境特征,以及林下优势种--紫耳箭竹的种群特征.结果表明: 随着落叶阔叶林→常绿落叶阔叶混交林→常绿阔叶林的演替,Shannon多样性指数、Simpson优势度指数和Pielou指数呈增加趋势,表明群落趋于稳定发展的状态;冠层结构特征也发生了显著改变,冠层开度和平均叶倾角减小,叶面积指数增加,冠层的消光能力增强,林下光照水平降低.上层林冠是造成林型郁闭的主要原因,其中冠层厚度和冠层面积是2个主要的影响因素.冠层结构与林下光照指标显著相关,对林下散射光的影响最大.冠层开度、林下光照条件均随着生长季的到来而下降,而叶面积指数呈现增长的趋势,峰值出现在6、7月;平均叶倾角在春季达到最大值,在夏季为最小值.紫耳箭竹的生长与冠层结构和光环境密切相关,其在光照适中的常绿落叶阔叶混交林中生长得最好,分株粗壮、密度大(29.69±1.68株·m^-2),地下茎拓展能力强;落叶阔叶林中的强光环境可能造成土壤水分缺失,从而对其生长产生影响;而在常绿阔叶林的低光环境下紫耳箭竹分株矮小,密度小(5.80±1.16株·m^-2),克隆扩展能力降低.在森林结构演变的过程中,冠层结构发生了明显的改变,显著影响林下光环境,过度的低光环境对紫耳箭竹种群的更新和发展有限制作用.     

Canopy manipulations of exotic Bitter Willow (Salix elaeagnos) forest for indigenous seedling recruitment: A pilot study

The invasive exotic tree species Bitter Willow (Salix elaeagnos; Salicaceae) has colonised areas of rank exotic grassland and has been found to contain indigenous seed, dispersed by frugivorous birds into the monospecific stands. This small pilot study examined whether indigenous seedlings that have germinated in the understorey of exotic Bitter Willow stands could be stimulated to establish through the creation of small‐scale canopy gaps. In Bitter Willow forest, four single Bitter Willow trees were poisoned to create canopy gaps. Light transmission and seedling regeneration of tree and shrub species were assessed beneath both the four manipulated and three comparable intact Bitter Willow canopies. Over 3 years, seedling height and density increased more beneath opened compared to intact Bitter Willow canopies. These results suggest that Bitter Willow can fill the roles of both a facilitative nurse and a perch tree. Larger‐scale canopy manipulation experiments of both Bitter Willow and other Salix species are needed to determine the full potential of canopy manipulations for forest restoration.     

GROWTH IN MEDEOLA VIRGINIANA CLONES. II. STOCHASTIC SIMULATION OF VEGETATIVE SPREAD

Medeola virginiana, a forest understory herb, has been hypothesized to minimize competition among ramets by positioning ramets regularly in space (Bell, 1974). We have constructed a computer model that simulates vegetative spread in Medeola by incorporating the variation in growth form found in natural populations. This stochastic model, based on data in Cook (1988), includes variation in the number and type of daughter ramets, the distance between parent and daughter ramets, and the angle of divergence between the lines of growth of the parent and daughter rhizomes. Over a wide range of initial ramet densities, initial degrees of ramet dispersion, and intensities of ramet mortality, our simulations indicate that the growth pattern of M. virginiana leads to aggregated distributions of ramets with overlapping leaf canopies. The observed aggregation of ramets may result from the tendency for directional but variable growth in clones. Such variable growth calls into question adaptive interpretations based on deterministic models that utilize the mean values of growth parameters but ignore the highly variable nature of clonal growth.     

Habitat selection in spatially heterogeneous environments: a test of foraging behaviour in the clonal submerged macrophyte Vallisneria spiralis

1. To test whether clonal macrophytes can select favourable habitats in heterogeneous environments, clonal fragments of the stoloniferous submerged macrophyte Vallisneria spiralis were subjected to conditions in which light intensity and substratum nutrients were patchily distributed. The allocation of biomass accumulation and ramet production of clones to the different patches was examined. 2. The proportion of both biomass and ramet number of clones allocated to rich patches was significantly higher than in poor patches. The greatest values of both clone and leaf biomass were produced in the heterogeneous light treatment, in which clones originally grew from light‐rich to light‐poor patches, while clones produced the most offspring ramets in the treatments with heterogeneous substratum nutrients. Similarly, root biomass had the highest values in nutrient‐rich patches when clones grew from nutrient‐rich to nutrient‐poor patches. 3. The quality of patches in which parent ramets established significantly influenced the foraging pattern. When previously established in rich patches, a higher proportion of biomass was allocated to rich patches, whereas a higher proportion of ramet number was allocated to rich patches when previously established in poor patches. 4. Results demonstrate that the clonal macrophyte V. spiralis can exhibit foraging in submerged heterogeneous environments: when established under resource‐rich conditions V. spiralis remained in favourable patches, whereas if established in adverse conditions it could escape by allocating more ramets to favourable patches.     

卧龙亚高山暗针叶林不同林冠环境下华西箭竹的克隆生长

该文调查了林下、中林窗、大林窗和林缘旷地等4种亚高山暗针叶林林冠环境下的华西箭竹(Fargesia nitida)分株种群,对其当年生立竹和母株的特征进行了比较研究,并探讨了母株年龄及大小对克隆生长的影响。主要研究结果如下:1)华西箭竹当年生立竹的分株密度以林下种群的最低,从林下→中林窗→大林窗,随林冠郁闭度的减小,华西箭竹基株当年生立竹的分株数逐渐增大。2)4种林冠环境下,当年生立竹和母株的分株高度、基径和生物量均有显著差异,且随林冠郁闭度的减小有递增的趋势(林下<中林窗<大林窗)。3)各林冠环境均以二龄母株产生的当年生立竹数量最大,不同林冠环境之间母株的平均年龄,以及处于同一林冠环境的各龄级母株产生的当年生立竹平均分株数和平均基径均无显著差异。4)不同林冠环境下,华西箭竹当年生立竹基径与一级母株、二级母株基径均呈正相关关系,但当年生立竹基径随一级母株增加的速度快于随二级母株的增加。5)当年生立竹的基径与地下茎直径呈显著的正相关;母株的基径与当年生立竹的地下茎直径呈正相关关系,而与其地下茎长度相关性不明显。     

Relationships between leaf nitrogen and limitations of photosynthesis in canopies of Solidago altissima

Vertical distribution patterns of light, leaf nitrogen, and leaf gas exchange through canopies of the clonal perennial Solidago altissima were studied in response to mowing and fertilizer application in a field experiment. Consistent with the distribution of light, average leaf nitrogen content followed a `smooth' exponential decline along the fertilized stands both in control and mown plots. The nitrogen profile along the unfertilized stands in mown plots, however, was `disrupted' by high-nitrogen leaves at the top of shorter ramets that only reached intermediate strata of the canopies. Hence, in these stands leaf nitrogen was significantly increased in short ramets compared with tall ramets for a given light environment, suggesting suboptimal stand structure but not necessarily suboptimal single-ramet architecture. However, at least under the climatic conditions observed during measurements, such disrupture had no substantial effect on stand productivity: model calculations showed that vertical distribution patterns of leaf nitrogen along ramets only marginally influenced the photosynthetic performance of ramets and stands. This is explained by the observed photosynthesis-nitrogen relationship: the rate of photosynthesis per unit amount of leaf nitrogen did not increase with leaf nitrogen content even under saturating light levels indicating that leaf photosynthesis was not nitrogen limited during the measurement periods. Nevertheless, our study indicates that consideration of how architecture(s) of adjacent individual plants interact could be essential for a better understanding of the trade-offs between individual and canopy characteristics for maximizing carbon gain. Such trade-offs may end up in a suboptimal canopy structure, which could not be predicted and understood by classical canopy optimization models.     

Gap-scale disturbance processes in secondary hardwood stands on the Cumberland Plateau,Tennessee, USA

Disturbance regimes in many temperate, old growth forests are characterized by gap-scale events. However, prior to a complex stage of development, canopy gaps may still serve as mechanisms for canopy tree replacement and stand structural changes associated with older forests. We investigated 40 canopy gaps in secondary hardwood stands on the Cumberland Plateau in Tennessee to analyze gap-scale disturbance processes in developing forests. Gap origin, age, land fraction, size, shape, orientation, and gap maker characteristics were documented to investigate gap formation mechanisms and physical gap attributes. We also quantified density and diversity within gaps, gap closure, and gap-phase replacement to examine the influence of localized disturbances on forest development. The majority of canopy gaps were single-treefall events caused by uprooted or snapped stems. The fraction of the forest in canopy gaps was within the range reported from old growth remnants throughout the region. However, gap size was smaller in the developing stands, indicating that secondary forests contain a higher density of smaller gaps. The majority of canopy gaps were projected to close by lateral crown expansion rather than height growth of subcanopy individuals. However, canopy gaps still provided a means for understory trees to recruit to larger size classes. This process may allow overtopped trees to reach intermediate positions, and eventually the canopy, after future disturbance events. Over half of the trees located in true gaps with intermediate crown classifications were Acer saccharum, A. rubrum, or Liriodendron tulipifera. Because the gaps were relatively small and close by lateral branch growth of perimeter trees, the most shade-tolerant A. saccharum has the greatest probability of becoming dominant in the canopy under the current disturbance regime. Half of the gap maker trees removed from the canopy were Quercus; however, Acer species are the most probable replacement trees. These data indicate that canopy gaps are important drivers of forest change prior to a complex stage of development. Even in relatively young forests, gaps provide the mechanisms for stands to develop a complex structure, and may be used to explain patterns of shifting species composition in secondary forests of eastern North America.     

Effect of waterlogging on regeneration in the dwarf birch (<Emphasis Type="Italic">Betula nana</Emphasis>)

The aim of this study was to determine the effect of waterlogging on annual regeneration in the dwarf birch (Betula nana L.), and to determine which demographic factors are responsible for the absence of B. nana in the waterlogged parts of a peat-bog. Growth parameters and demographic data were collected from 2001 to 2003 at four sites in the Linje peat bog in the Chełmno Lake District in northern Poland. Three of the sites represented waterlogged open bog communities. The fourth site served as the reference. The density of first-year ramets, annual shoot growth, and the proportion of flowering shoots was significantly lower at the waterlogged sites than at the reference site. No seedlings were found during the course of the study, even though flowering and fruit production took place at the reference site. The flowering shoots were mainly between nine and thirteen years old. Ramets in this age range were found only at the reference site due to high mortality in mature ramets at the waterlogged sites. Mean ramet height was significantly lower at the waterlogged sites than at the reference site, and was strongly correlated with mean ramet age. The distribution of B. nana in peat bogs is limited primarily by two demographic factors: the high mortality rate in older ramets, and the low rate of vegetative reproduction. Another contribution factor is that the proportion of older fertile ramets is low. The effects of fluctuations in the water table on distribution of B. nana are also discussed.