Stochasticity of species assemblage of canopy trees and understorey plants in a temperate secondary forest created by major disturbances

Detrended canonical correspondence analysis (DCCA) was applied to explore the species assemblage of plants in a temperate secondary forest that was created by major disturbances. The DCCA showed vague relationships between species dominance and environmental factors for canopy tree species even when rare species were excluded from the analysis. For the highest dominant species of the understorey, the scores of the first axis, which correlated with the species richness of overstorey trees, decreased. This fact showed that the assemblage of canopy trees affects, through the differences in leaf phenology or leaf characteristics of canopy trees, the life history of dominant understorey plants. The studys results suggest that competition colonization might be more important for canopy trees during the developing stage of the forest if the disturbance occurs on a large scale, and that colonization from the local species pool determines species assemblage on a landscape scale.     

Importance of seed size for the establishment of seedlings of five deciduous broad-leaved tree species

Effects of seed size and phenology on the establishment of five deciduous broad-leaved tree species were examined in deciduous woodland. Treatments included absence and presence of litter in the forest understory, a small gap, and a large gap. Seedling emergence of large-seeded speciesQuercus mongolica var.grosseserrata andAcer mono was not reduced by accumulation of litter in the forest understory, but was promoted in the large gap where litter was less. Seedling emergence of small-seeded species,Alnus hirsuta, Cercidiphyllum japonicum andBetula platyphylla var.japonica, was reduced by the litter in almost all of the sites. Seedlings of large-seeded species avoid shade stress phenologically by unfolding all of their large leaves in a short period before canopy closure in the forest understory. These species had little mortality after seedling emergence. In contrast, small-seeded species have a longer duration of leaf emergence, shorter leaf longevity, and rapid leaf turnover in all the sites. These seedlings attained similar height to those of the large-seeded species at the end of the second year in the large gap, but survival and height growth rate decreased after canopy closure in the forest understory. We suggest that the importance of seed size in determining seedling establishment largely depends on the relationships between seasonal changes of environmental conditions and phenological traits of seedlings, which are related to seed size.Abbreviations Ah Alnus hirsuta - Am Acer mono - Cj Cercidiphyllum japonicum - Bp Betula platyphylla var.japonica - Qm Quercus mongolica var.grosseserrata     

Leaf display and photosynthesis of tree seedlings in a cool-temperate deciduous broadleaf forest understorey

To examine a possible convergence in leaf photosynthetic characteristics and leaf display responses to light environment in seedlings of three canopy and two shrub tree species in understorey of cool-temperate deciduous broadleaf forest, relationships between light environment, leaf orientation and leaf light-photosynthetic response were measured. Light capture of the seedlings (17-24 individuals with 2-12 leaves for each species) was assessed with a three dimensional geometric modeling program Y-plant. Leaf photosynthetic characteristics of the five species were found to have acclimated to the understorey light environment, i.e., low light compensation point and high apparent quantum yield. In addition, light-saturated photosynthetic rates were higher in seedlings inhabiting microsites with higher light availability. Efficiencies of light capture and carbon gain of the leaf display were evaluated by simulating the directionalities of light capture and daily photosynthesis for each seedling using hemispherical canopy photography. The results showed that most of the seedlings orientated their leaves in a way to increase the daily photosynthesis during the direct light periods (sunflecks) rather than maximize daily photosynthesis by diffuse light. Simulations also showed that daily photosynthesis would increase only 10% of that on actual leaf display when the leaves orientated to maximize the diffuse light interception. Simulations in which leaf orientations were varied showed that when the leaf display fully maximized direct light interception, the time that leaves were exposed to excessive photon flux density of >800 mumol photons m(-2) s(-1) were doubled. The understorey seedlings studied responded to the given light environments in a way to maximize the efficiency of acquisition and use of light during their short (approximately 3 month) seasonal growth period.     

Temperature effects on forest herbs assessed by warming and transplant experiments along a latitudinal gradient

Slow‐colonizing forest understorey plants are probably not able to rapidly adjust their distribution range following large‐scale climate change. Therefore, the acclimation potential to climate change within their actual occupied habitats will likely be key for their short‐ and long‐term persistence. We combined transplant experiments along a latitudinal gradient with open‐top chambers to assess the effects of temperature on phenology, growth and reproductive performance of multiple populations of slow‐colonizing understorey plants, using the spring flowering geophytic forb Anemone nemorosa and the early summer flowering grass Milium effusum as study species. In both species, emergence time and start of flowering clearly advanced with increasing temperatures. Vegetative growth (plant height, aboveground biomass) and reproductive success (seed mass, seed germination and germinable seed output) of A. nemorosa benefited from higher temperatures. Climate warming may thus increase future competitive ability and colonization rates of this species. Apart from the effects on phenology, growth and reproductive performance of M. effusum generally decreased when transplanted southwards (e.g., plant size and number of individuals decreased towards the south) and was probably more limited by light availability in the south. Specific leaf area of both species increased when transplanted southwards, but decreased with open‐top chamber installation in A. nemorosa. In general, individuals of both species transplanted at the home site performed best, suggesting local adaptation. We conclude that contrasting understorey plants may display divergent plasticity in response to changing temperatures which may alter future understorey community dynamics.     

我国温带山地森林48种常见树种叶片重量-出叶强度的关系

叶片是植物的主要光合器官, 其质量与数量的权衡关系体现植物对环境的适应策略。在全球气候变化的背景下, 研究叶片质量与数量关系有助于理解植物对环境变化的响应趋势。该研究应用标准化主轴回归方法, 探讨了我国温带山地森林中48个常见树种的单叶干重与出叶强度的权衡关系。结果表明, 所有物种以及落叶阔叶林、常绿和落叶阔叶树种、单叶以及亚冠层阔叶树种的单叶干重与出叶强度表现为异速生长关系; 针叶林、针阔混交林、常绿及落叶针叶树种、复叶以及冠层阔叶树种则表现为等速生长关系。研究结果表明, 叶大小和出叶强度并无恒定的权衡关系。     

Tree fern trunks facilitate seedling regeneration in a productive lowland temperate rain forest

Seedling regeneration on forest floors is often impaired by competition with established plants. In some lowland temperate rain forests, tree fern trunks provide safe sites on which tree species establish, and grow large enough to take root in the ground and persist. Here we explore the competitive and facilitative effects of two tree fern species, Cyathea smithii and Dicksonia squarrosa, on the epiphytic regeneration of tree species in nutrient-rich alluvial forests in New Zealand. The difficulties that seedlings have in establishing on vertical tree fern trunks were indicated by the following observations. First, seedling abundance was greatest on the oldest sections of tree fern trunks, near the base, suggesting that trunks gradually recruited more and more seedlings over time, but many sections of trunk were devoid of seedlings, indicating the difficulty of establishment on a vertical surface. Second, most seedlings were from small-seeded species, presumably because smaller seeds can easily lodge on tree fern trunks. Deer browsing damage was observed on 73% of epiphytic seedlings growing within 2 m of the ground, whereas few seedlings above that height were browsed. This suggests that tree ferns provide refugia from introduced deer, and may slow the decline in population size of deer-preferred species. We reasoned that tree ferns would compete with epiphytic seedlings for light, because below the tree fern canopy photosynthetically active radiation (PAR) was about 1% of above-canopy PAR. Frond removal almost tripled %PAR on the forest floor, leading to a significant increase in the height growth rate (HGR) of seedlings planted on the forest floor, but having no effects on the HGRs of epiphytic seedlings. Our study shows evidence of direct facilitative interactions by tree ferns during seedling establishment in plant communities associated with nutrient-rich soils.     

Trade-offs between seedling growth and survival in deciduous broadleaved trees in a temperate forest

BACKGROUND AND AIMS: In spatially heterogeneous environments, a trade-off between seedling survival and relative growth rate may promote the coexistence of plant species. In temperate forests, however, little support for this hypothesis has been found under field conditions, as compared with shade-house experiments. Performance trade-offs were examined over a large resource gradient in a temperate hardwood forest. METHODS: The relationship between seedling survival and seedling relative growth rate in mass (RGR(M)) or height (RGR(H)) was examined at three levels of canopy cover (forest understorey, FU; small gap, SG; and large gap, LG) and at two microsites within each level of canopy cover (presence or absence of leaf litter) for five deciduous broad-leaved tree species with different seed sizes. KEY RESULTS: Within each species, both RGR(M) and RGR(H) usually increased with increasing light levels (in the order FU < SG < LG), whereas little difference was observed based on the presence or absence of litter. Seedling survival in FU was negatively correlated with both RGR(M) and RGR(H) in both LG and SG. The trade-off between high-light growth and low-light survival was more evident in the relationship with LG as compared with SG. An intraspecific trade-off between survival and RGR was observed along environmental gradients in Acer mono, whereas seedlings of Betula platyphylla var. japonica survived and grew better in LG. CONCLUSIONS: The results presented here strongly support the idea of light gradient partitioning (i.e. species coexistence) in spatially heterogeneous light environments in temperate forests, and that further species diversity would be promoted by increased spatial heterogeneity. The intraspecific trade-off between survival and RGR in Acer suggests that it has broad habitat requirements, whereas Betula has narrow habitat requirements and specializes in high-light environments.     

Tree fruiting phenology in Kalinzu Forest, Uganda

The spatial and temporal variations in species composition and abundance of trees that fruited in mechanically logged, intensively pit‐sawn and essentially undisturbed forest areas in Kalinzu Forest (0°17′S, 0°30′S and 30°00′, 30°07′E) were assessed. The duration and frequency of fruiting episodes of selected pioneer, understorey and canopy tree species were also determined. These phenology aspects were monitored for 18 months (between February 1997 and July 1998) in 99 plots, each 20 m × 20 m in size. Eighty species consisting of 1489 trees fruited during the study. Most of the species were recorded in the undisturbed forest area and least in the mechanically logged areas. Monthly number of individuals and species that fruited in each forest condition were significantly related to monthly rainfall. Two pioneer species (Musanga leo‐errerae and Trema orientalis), one sub‐canopy (Funtumia africana) and canopy (Parinari excelsa) species had extended fruiting phases (>5 months). Two understorey species (Oxyancius speciosus and Tabernaemontana spp.) had an extended phase while the third (Teclea nobilis) had a short (≤5 months) fruiting phase. Of the selected species, only Strombosia scheffleri (a sub‐canopy species) and P. excelsa had more than one fruiting episode.     

南京地区落叶栎林主要木本植物的展叶动态研究

 植物的展叶过程是由自身遗传因子决定的,同时又受到多种生态因子的调节,反映了植物的生活史对策和群落物种多样性的维持机制。在2001和2002年的3～6月间,不定期记录了南京地区三个落叶栎(Quercus spp.)林中主要木本植物的展叶情况,包括被标记标准枝的叶数、叶的长度、宽度、叶面积、叶干重等参数。结果表明在所调查的落叶栎林中,林冠层物种的成熟叶面积和单位叶面积干重都显著大于林下层物种;最早展叶的物种为林下层物种,但林冠层展叶顺序与林下层无显著差异。叶面积越大、单位叶面积干重越小的物种展叶越晚;林冠层物种展叶较林下层快,物种成熟叶面积越大,展叶速率越大。最后对展叶顺序和展叶速度的生态学意义作了讨论。     

猫儿山不同海拔植物群落树木构型差异及其影响因子

通过对猫儿山海拔梯度上常绿阔叶林(低海拔,1100 m)、常绿落叶阔叶混交林(中海拔,1500 m)和常绿针阔叶混交林(高海拔,1900 m)3种典型植物群落中乔木层植物构型性状以及环境因子的测定,分析乔木层植物构型性状在3个群落间的变异规律及其影响因素.结果表明:随海拔升高,乔木层树冠面积、45 cm基径、胸径和叶片聚集度持续增加,树高、枝下高和树冠厚度先增加后减小;枝条伸展方向表现为在低海拔群落中水平枝条比例最大,高海拔群落次之,中海拔群落最小;中海拔群落中乔木层植物构型性状之间的相关性更强.冗余分析表明,土壤有机质和总辐射是乔木层植物构型性状变异的主要环境影响因素,它们分别解释了构型性状39.6%和23.9%的变异.土壤有机质对树冠面积和枝下高影响较大,总辐射对胸径和45 cm基径影响较大,且都呈正相关.猫儿山乔木层植物在不同海拔群落间存在构型分异,影响乔木层植物构型变化的主要环境驱动力是土壤有机质和总辐射.     

Overstorey Control of Understorey Species Composition in a Near-natural Temperate Broadleaved Forest in Denmark

Little is known about the importance of the forest overstorey relative to other factors in controlling the spatial variability in understorey species composition in near-natural temperate broadleaved forests. We addressed this question for the 19 ha ancient forest Suserup Skov (55°22′ N, 11°34′ E) in Denmark, one of the few old-growth temperate broadleaved forest remnants in north-western Europe, by inventorying understorey species composition and environmental conditions in 163 100 m² plots. We use unconstrained and constrained ordinations, variation partitioning, and Indicator Species Analysis to provide a quantitative assessment of the importance of the forest overstorey in controlling understorey species composition. Comparison of the gradients extracted by unconstrained and constrained ordinations showed that the main gradients in understorey species composition in our old-growth temperate broadleaved forest remnant are not caused by variability in the forest overstorey, but are related to topography and soil, edge effects, and unknown broad-scale factors. Nevertheless, overstorey-related variables uniquely accounted for 15% of the total explained variation in understorey species composition, with the pure overstorey-related (R_po), topography and soil (R_pt), edge and anthropogenic disturbance effects (R_pa), and spatial (R_ps) variation fractions being of equal magnitude. The forward variable selection showed that among the overstorey-related variables understorey light availability and to a lesser extent vertical forest structure were most important for understorey species composition. No unique influence of overstorey tree species identity could be documented. There were many indicator species for high understorey light levels and canopy gap centres, but none for medium or low light or closed canopy. Hence, no understorey species behaved as obligate shade plants. Our study shows that, the forest overstorey has a weak control of understorey species composition in near-natural broadleaved forest, in contrast to results from natural and managed forests comprising both conifer and broadleaved species. Nevertheless, >20% of the understorey species found were indicators of high light conditions or canopy openings. Hence, variability in canopy structure and understorey light availability is important for maintaining understorey species diversity.     

Development of Tree Regeneration in Fern-dominated Forest Understories After Reduction of Deer Browsing

Dennstaedtia punctilobula (hay‐scented fern) can act as a native invasive species in forests in eastern North America where prolonged deer browsing occurs in stands with partially open overstory canopies. Ferns dominate the understory with a 60‐cm tall canopy, with little regeneration of native tree species. It has been hypothesized that, once established, ferns may continue to inhibit tree regeneration after deer browsing has been reduced. To test this hypothesis, we documented the pattern of recovery of the tree seedling understory in plantations of Pinus strobus (white pine) and Pinus resinosa (red pine) on the Quabbin Reservation watershed protection forest in central Massachusetts, where after 40 years of intensive deer browsing the deer herd was rapidly reduced through controlled hunting. Dense fern understories occur on nearly 4,000 ha of the predominantly oak–pine forest. Three years after deer herd reduction, stands with the highest density fern cover (77% of plots with>90% cover) had significantly fewer seedlings at least 30 cm in height, compared with stands with lower fern density, and those seedlings consisted almost entirely of Betula lenta (black birch) and white pine. Height growth analysis showed that black birch and white pine grew above the height of the fern canopy in 3 and 6 years, respectively. In contrast, two common species, Fraxinus americana (white ash) and Quercus rubra (red oak), grew beneath the dense fern cover for 5 years with height growth less than 5 cm/yr after the first year. A study of spring phenology indicated that the ability of black birch to grow through the fern canopy might have been due to its early leaf development in spring before the fern canopy was formed, in contrast to oak and ash with delayed leaf development. Thus, the ferns showed differential interference among species with seedling development after reduction of deer browse.     

Tree species distribution in canopy gaps and mature forest in an area of cloud forest of the Ibitipoca Range,south-eastern Brazil

The tree community of both canopy gaps and mature forest was surveyed in a 5 ha plot of cloud forest in the Ibitipoca Range, south-eastern Brazil, aiming at: (a) comparing the tree community structure of canopy gaps with that of three strata of the mature forest, and (b) relating the tree community structure of canopy gaps with environmental and biotic variables. All saplings of canopy trees with 1–5 m of height established in 31 canopy gaps found within the plot were identified and measured. Mature forest trees with dbh 3 cm were sampled in four 40×40 quadrats laid on the four soil sites recognised in the local soil catena. All surveyed trees were identified, measured and distributed into three forest strata: understorey (<5 m of height), sub-canopy (5.1–15 m) and canopy (15.1–30 m). The following variables were obtained for each gap: mode of formation, age, soil site, slope grade, size, canopy openness and abundance of bamboos and lianas. A detrended correspondence analysis indicated that the tree community structure of gaps in all soil sites was more similar to that of the mature forest understorey, suggesting that the bank of immatures plays an important role in rebuilding the forest canopy and that gap phases may be important for understorey shade-tolerant species. There was evidence of gap-dependence for establishment for only one canopy tree species. Both canonical correspondence analysis and correlation analysis demonstrated for a number of tree species that the distribution of their saplings in canopy gaps was significantly correlated with two variables: soil site and canopy openness. The future forest structure at each gap is probably highly influenced by both the present structure of the adjacent mature forest and the gap creation event.     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

不同邻体大头茶幼苗的适应特征

 在亚热带常绿阔叶林自然林窗下,以大头茶幼苗为基株,分别以马尾松、川灰木和大头茶幼苗为邻体。研究不同邻体如何改变可利用资源,影响大头茶幼苗基株生长动态和形态、树冠结构和生物量效应,并探讨影响基株生长的邻体的主导因素。结果表明,不同邻体的生长和截光效应差异明显;基株生长特征与邻体的树高、叶生物量、总生物量和树冠距地表10cm处截光量相关极小,却与邻体树冠距地表30cm处截光量相关极大,故影响基株生长的邻体主导因素是遮挡基株树冠的邻体的可利用光资源;基株生长动态,如树高、叶数、叶面积和叶大小,树冠结构和生物量显著受到不同邻体的影响,并且基株树高及增长量、叶面积、冠幅和生物量均为大头茶单株(GNO)>大头茶+川灰木(GNS)>大头茶+马尾松(GNP)>大头茶+大头茶(GNG)。大头茶幼苗种内竞争明显强于种间竞争。     

Tree seed germination and seedling establishment in treefall gaps and understorey in a subtropical forest of northeast India

Abstract Seed germination, and survival and growth of seedlings of four dominant tree species, Quercus dealbata, Quercus griffithii, Quercus glauca and Schima khasiana were studied in the treefall gaps and forest understorey of an undisturbed mature-phase humid subtropical broadleaved forest in northeast India. Three important microenvironmental factors namely photosynthetically active radiation (PAR), soil moisture and litter depth, were also measured in the forest understorey and gaps and correlated with seedling mortality. Seed germination of S. khasiana was significantly higher in the treefall gaps than in the understorey; among the tree species studied, it had the highest germination. Quercus seedlings were abundant in the understorey and small gaps, while S. khasiana seedlings were more numerous in the large gaps. The survivorship curves for the seedling populations revealed that the three Quercus species survived better in the understorey, while S. khasiana did so in the gaps. PAR and soil moisture were positively correlated with tree seedling mortality, which occurred mainly during the winter months. The Quercus seedlings grew better in the forest understorey and small gaps and S. khasiana seedlings in the large gaps. The differential performance of the tree seedlings to the conditions prevailing in the understorey and gaps of two sizes indicates that different species were adapted to different light environments depending upon their optimum requirements. This could be an effective mechanism for promoting species coexistence in the forest community.     

Crown structure and leaf area of the understorey species Prunus serotina

A detailed biometrical study of the exotic understorey invader Prunus serotina (Ehrh.) was carried out in a mixed coniferous forest stand in northern Belgium. Based on detailed destructive measurements of eight selected model trees, allometric relations of tree height, crown projected area, woody and leaf dry mass and leaf area on tree diameter at breast height (DBH) were derived. The scaling-up procedure from the tree to the stand level was done using the frequency distribution of DBH obtained at the selected experimental plot. The vertical and radial distributions of the tree foliage were estimated by the “cloud” technique. The vertical profile of leaf area showed a bimodal distribution pattern with maxima at heights of 4 and 6 m above the ground. The leaf area index (LAI) of the understorey Prunus serotina as estimated by the described up-scaling procedure (5.1) was significantly higher than the LAI (2.6) as measured by a plant canopy analyser and was also significantly higher than the LAI of the overstorey species Scots pine (1.5–3.0). The LAI of a neighbouring Rhododendron understorey reached only 1.25. This study emphasises the importance of an exotic understorey species in the total leaf area of mixed coniferous forests which might have important implications for the energy and mass exchanges of the entire forest.     

Leaf traits determine the growth-survival trade-off across rain forest tree species

A dominant hypothesis explaining tree species coexistence in tropical forest is that trade-offs in characters allow species to adapt to different light environments, but tests for this hypothesis are scarce. This study is the first that uses a theoretical plant growth model to link leaf trade-offs to whole-plant performances and to differential performances across species in different light environments. Using data of 50 sympatric tree species from a Bolivian rain forest, we observed that specific leaf area and photosynthetic capacity codetermined interspecific height growth variation in a forest gap; that leaf survival rate determined the variation in plant survival rate under a closed canopy; that predicted height growth and plant survival rate matched field observations; and that fast-growing species had low survival rates for both field and predicted values. These results show how leaf trade-offs influence differential tree performance and tree species' coexistence in a heterogeneous light environment.     

降水变化和种间竞争对红松和蒙古栎幼苗生长的影响

针对全球变暖导致的降水格局的变化,选择长白山红松针阔叶混交林主要树种蒙古栎和红松幼苗为研究对象,在野外自然条件下人工模拟增、减水30%对单种和混种的针阔叶树种形态、生长和生物量分配的影响. 结果表明：对于蒙古栎幼苗,与单种处理相比,混种显著增加了其冠幅和主根长,与对照处理相比,减水处理显著提高了其茎质比、降低了其主根长; 对于红松幼苗,与单种处理相比,混种显著减少了其基径、树高、叶片数和根、茎、叶质量及总干质量,与对照处理相比,减水处理显著降低了其主根长、叶片数、叶质量和总干质量以及叶质比,同时显著提高了茎质比. 增水处理对二者的影响均不显著. 在树木生长初期,种间竞争和降水格局变化对蒙古栎和红松幼苗形态和生长均产生显著影响,对红松幼苗的影响更大.