Factors influencing establishment by direct seeding of indigenous tree species in typical plantations and shrubland in South China

Seed limitation of desired indigenous species has constrained the succession of established plantations from an early successional stage, to more natural forests in South China. Because of its low cost and time-saving advantages, direct seeding has been considered a useful method in forest regeneration. To improve our understanding of whether and how direct seeding can lead to the establishment of indigenous tree species, we carried out an experiment in four typical plantations (eucalyptus, mixed-native, mixed-legume, mixed-conifer) and a shrubland in South China. We also tested the effect of understory vegetation and litter removal on seedling establishment and growth. Seeds of three indigenous tree species (Castanopsis chinensis, Cryptocarya chinensis, and Psychotria rubra) were sown with understory vegetation and litter either retained or removed. Seedling emergence, survival, and growth were recorded for 21 months. We found that conditions that favored seedling emergence often differed from those that favored seedling survival. While the removal of understory vegetation and litter did not influence the final seedling establishments for all the three introduced species, growth of Psychotria rubra was influenced at several sites. Seed mass was positively correlated with seedling emergence and subsequent growth. The large-seeded Castanopsis chinensis showed higher emergence and growth than the intermediate-seeded Cryptocarya chinensis or the small-seeded Psychotria rubra. Seedlings of Castanopsis chinensis grew better in plant communities with high understory light penetration and high soil exchangeable potassium. Growth of P. rubra seedlings decreased significantly with soil hydrolyzed nitrogen. In conclusion, direct seeding can be useful for the establishment of indigenous tree species into plantations and shrubland. Optimizing early seedling establishment and growth will require the selection of suitable native species for improving seedling establishment in different plantations. Additionally, understory vegetation and litter and fertilizer should be properly managed to enhance the establishment of certain indigenous tree species.     

Tree recruitment in gaps of various size,clearcuts and undisturbed mixed forest of interior British Columbia,Canada

Tree seedling recruitment was monitored after various types of logging in mixed conifer and deciduous forests of northern British Columbia, Canada. Predicting tree seedling recruitment after disturbance is fundamental to understanding forest dynamics and succession and is vital for forest management purposes. Seedling recruitment success in multi-species northern latitude forests varied as a function of mature tree canopy cover, gap size and position in a gap. Recruitment was abundant within canopy gaps across a wide range of gap sizes (20–5000 m²), but recruit numbers dropped off rapidly under the closed forest canopy and in the open conditions of clearcuts. Inside canopy gaps, recruitment was similar by gap position in small gaps (<300 m²) but, in these northern latitude forests, exhibited a trend of increasing density from the sunny north to shady south end of larger gaps. This was true for all tree species regardless of their shade tolerance ranking. There was no evidence of gap partitioning by any of the tree species during the regeneration phase suggesting that adaptation to the subtleties of gap size during early recruitment are not well developed in these tree species. Favorable locations for emergence and early establishment of germinants were less favorable for growth and survival of established seedlings, i.e. the regeneration niches in these forests were discordant. Tree abundance and species diversity appears to be controlled more by differentiation among growth and survival niches than by the regeneration niches. From the perspective of forest management, abundant natural regeneration of all the dominant tree species of these mixed-species forests can be obtained after partial cutting.     

Tree mortality and habitat shifts in the regeneration trajectory underneath canopy of an old-growth subalpine forest

To better understand tree regeneration trajectories and the resultant coexistence of Abies with co-dominants, Picea jezoensis var. hondoensis, Tsuga diversifolia and Betula ermanii, in an old-growth subalpine forest, we investigated spatial mortality patterns during the regeneration of Abies mariesii and A. veitchii, which are abundant in the understory reflecting their shade tolerance. Regeneration of these Abies spp. from shaded understory to canopy status is affected by other canopy co-dominants. Snags of understory Abies spp. were common, suggesting that the primary mortality agent is suppression by the overstory. Although live, small Abies trees in the understory were positively associated with a Picea canopy, the long-term survival was reduced among Abies trees close to the canopy, suggesting that shading by large Picea in the overstory negatively affects understory Abies plants. The existence of shade-intolerant canopy co-dominants such as Picea and also Tsuga, which are larger and longer lived than the shade-tolerant Abies, may play an important role in preventing the Abies spp. from competitively displacing these other tree species, which are much rarer in the understory, though common in the canopy. Moreover, in spite of the fact that Betula canopies fostered recruitment and growth of Abies saplings, Abies showed no association with Betula canopy and their survival at later-stage was rather reduced near or beneath Betula canopies at the subsequent understory small tree stage. Based on spatially significant events related to tree death, this study detected such “habitat shifts” in the trajectory of tree regeneration. Accordingly, it can be concluded that careful consideration of the regeneration habitat is required for a fuller understanding of ecological processes in spatially complex old-growth forest systems.     

Canopy effects on vegetation caused by harvesting and regeneration treatments

This paper reviews the primary effects of canopy on understorey vegetation to provide insight into the management of the canopy space to benefit tree regeneration. Site conditions, like nutrient and water availability, overstorey conditions, e.g. tree species, and canopy density, are important determinants of ground vegetation. An investigation of canopy effects is concerned about how the canopy trees modify site conditions. As canopy density may be deliberately modified during regeneration treatments, the effect of canopy density on individual species in the herbaceous layer and tree regeneration is important. This autecological perspective focuses on the successional traits of species to help understand species differences in fecundity, survival, density and growth. From a synecological perspective, the importance of successional traits for the outcome of competition between species arising from differences in canopy densities is highlighted. This review shows that moderately dense canopies may favour tree regeneration over aggressive shade-intolerant graminoids or forbs. This is particularly true for shade-tolerant and intermediate shade-tolerant tree species. To better understand and utilise this phenomenon, research should try to identify and isolate different canopy effects.     

Floristic and structural recovery of a laurel forest community after clear-cutting: A 60 years chronosequence on La Palma (Canary Islands)

We analyzed a post-clearcut chronosequence (0.5 to 60 years after harvesting) in the laurel forest of La Palma island (Canarian Archipelago) to determine the recovery of the stands with respect to species composition, richness, life strategies and structural parameters of the canopy. Multivariate analysis showed that exotic species, as well as annual ruderal species were confined to early-successional stages, while native perennials, typical of laurel forests, dominated the late-successional stages. Total species richness decreased significantly with time after clear-cutting. The relative fast recovery of understory native species may be due to low forest floor disturbance during harvesting. Shade-intolerant pioneer, pioneer-remnant and shade-tolerant late-successional species were the main life strategies of native tree species. Most structural parameters showed a continuous and monotonic increase (basal area, biomass) or decrease (density, percentage of photosynthetic biomass) during succession. Once clear-cutting, here performed with an interval of 8 years, is abandoned, the recovery of the laurel forest seems possible due to careful logging that protects the soil and a rapid asexual regeneration of native tree species, revealing this to be a sustainable management practice.     

Effect of clear-cutting on the understory vegetation,soil and diversity of litter beetles in scots pine-dominated forest

Currently, the aim of modern forest management is not only timber production but also the protection of biodiversity. The initial effects of clear-cut logging on forest understory vegetation, soil and the diversity of litter beetles have been studied. We examined unstudied effects of clear-cutting disturbance on understory vascular plant species abundance, forest soils, and understory vegetation nutrients as well as beetle diversity one year after clear-cutting. Substantial changes in the prominence values and above-ground mass of forest-related vascular plant and moss species were detected. Clear-cuttings resulted in fast appearance of new light-loving plant species. The significant decrease in understory plant biomass influenced the reduction of nutrient pools in clear-cut areas after one year. The clear-cut logging negatively influenced the richness of species and the number of individuals of stenobiontic forest-related beetles in the forest litter. However, forest clear-cuttings were beneficial for eurytopic and open land species.     

Secondary edge effects in regenerating forest landscapes: vegetation and microclimate patterns and their implications for management and conservation

Forest regrowth is expected to gradually mitigate edge effects in forest landscapes fragmented by timber harvest, but our understanding of edge effect persistence and dynamics over time is still incomplete. Our main objective was to take a critical look at the role of forest regrowth in mitigating the initial edge effects on microclimate and understory vegetation in northern hardwood forests of the eastern United States. We compared canopy closure, hourly air temperature, soil moisture, and understory vegetation at increasing distances from forest edges (0, 5, 10, 20, and 30 m) along twelve transects placed across new and older forest edges (3–4 or 16–19 years old) created by forest harvest. Open, new forest edges exhibited pronounced edge effects on microclimate and shade-intolerant plants, but these were almost completely moderated by forest regrowth on the cleared side of older edges where dense young forest developed with a new canopy comparable in cover to adjacent mature forest. There were no initial edge effects on shade-tolerant vegetation across new forest edges, but the shade-tolerant vegetation declined in mature forest near old forest edges adjacent to dense young forest that supported only sparse understory vegetation. These delayed secondary edge effects of young dense forests on adjacent mature forests have not been previously documented and they should be more explicitly included in forest management considerations. We suggest an integrated system for managing and mitigating both the immediate primary and delayed secondary edge effects in those working forest landscapes where biodiversity conservation is of high priority.     

Tree regeneration in gap-understory mosaics in a subtropical Shorea robusta(Sal) forest

Spatial variation in tree-regeneration density is attributed to the specialization of tree species to light availability for germination and growth. Light availability,in turn, varies across the gap-understorey mosaic. Canopy gaps provide an important habitat for the regeneration of tree species that would otherwise be suppressed in the understory. In subtropical forests, there is still a knowledge-gap relating to how canopy disturbances influence tree-regeneration patterns at local scale, and if they disproportionately favor regeneration of certain species. We aim to analyze whether canopy gaps promote tree regeneration, and tree species are specialized to gaps or understory for germination and growth. We sampled vegetation in 128 plots(0.01 ha), equally distributed in gaps and below canopy, in two subtropical Shorea robusta Gaertn.(Sal) forests in Nepal, recording the number of tree seedlings and saplings in each plot. We compared the regeneration density of seedlings and saplings separately between gaps and the understorey. The mean densities of seedlings and saplings were higher in the gaps at both sites;although there was no difference in the seedling density of the majority of the species between the habitats. No species were confined to either gap or understorey at the seedling stage. We conclude that gaps are not critical for the germination of tree species in Sal forests but these are an important habitat for enabling seedlings to survive into saplings. The classification of trees into regeneration guilds mainly based on germination does not apply to the majority of tree species in subtropical Sal forests. Our results reaffirm that gap creation promotes tree regeneration by favouring seedling survival and growth and can influence forest management for conservation, as well as for plantations.     

Heavy thinning in hinoki plantations in Shikoku (southwestern Japan) has limited effects on recruitment of seedlings of other tree species

Heavy thinning may provide a feasible means to convert single-species coniferous plantations to broadleaved or mixed forests. To assess this possibility, we monitored tree seedling dynamics of other tree species, the understory microenvironment, and seed availability following heavy thinning (50 % by volume) in two hinoki (Chamaecyparis obtusa) plantations in Shikoku, southwestern Japan. Conventional (35 % by volume) and row thinning (50 % by volume) treatments were also applied at one of the sites. Diffuse transmittance in the understory dramatically increased after all thinning treatments then rapidly decreased, except after row thinning. Heavy thinning accelerated recruitment of small seedlings (with stems <30 cm high) at one of the sites. However, the densities of these seedlings declined sharply, and increases in the densities of larger seedlings recorded in subsequent surveys were very small. Furthermore, at the other site, recruitment of small seedlings under heavy thinning was equivalent to that under conventional thinning and lower than that under row thinning. Abundant species in the recruited seedlings were pioneer or subcanopy species found in the soil seed bank. Low seed rain density at both sites appeared to be responsible for the poor recruitment of canopy tree species. Our results suggest that heavy thinning had limited effects on the seedling recruitment of other tree species in these hinoki plantations due to insufficient seed rain and rapid canopy closure.     

Establishment and early growth of introduced indigenous tree species in typical plantations and shrubland in South China

Unsuccessful colonization by indigenous tree species into established plantations has retarded the succession of artificial plantations to more natural, secondary forests in South China. To understand how to improve colonization by seedlings of indigenous species, we determined how performance of indigenous seedlings is affected by seedling species (the shade-intolerant Castanopsis chinensis, the moderately shade-intolerant Michelia chapensis, and the shade-tolerant Psychotria rubra), the site into which the seedlings were transplanted (a mixed-legume plantation, a eucalyptus plantation, a mixed-native plantation, a mixed-conifer plantation, and a shrubland), and site preparation (removal or retention of understory vegetation and litter). Seedling survival and growth were generally increased by removal of understory vegetation and litter. C. chinensis and M. chapensis grew better in the mixed-legume and mixed-conifer plantations, while P. rubra grew better in mixed-native and mixed-conifer plantations. Responses of the transplanted seedlings to environmental factors were species specific. The effects of light on seedling survival and growth were correlated with the shade tolerance of the species. Soil moisture was important; it was positively correlated with survival but negatively correlated with growth of C. chinensis seedlings. Growth of C. chinensis and M. chapensis was positively correlated with soil potassium, while growth of P. rubra was positively correlated with soil organic matter but negatively correlated with soil hydrolyzed nitrogen. These findings suggest that we should select suitable native species under the different plantations before improvement of plantations. Light and soil moisture are most important environmental factors for the selection of species specific. Site preparation and fertilizer are needed during the improvement of those plantations.     

Tree regeneration and plant species diversity responses to vegetation control following a major windthrow in mixed broadleaved stands

By increasing resource availability, canopy opening enhances tree recruitment as well as the development of neighbouring vegetation. The proliferation of early successional and highly competitive vegetation may have dramatic consequences on seedling establishment. However, differences in competitive abilities have been shown among the plant growth forms commonly encountered in forests. We may thus expect that vegetation management leading to control of different plant growth forms would have different consequences on tree seedling growth and development. To test this hypothesis, we analysed the effects of an intensity gradient of four vegetation control treatments (untreated, coppice control, coppice and non-tree plant control, and coppice, non-tree and pioneer (tree) plant control) on plant species richness and natural tree regeneration in three post-storm sites. Higher plant species richness and a better natural tree regeneration were observed in the more intensive treatments that significantly improved the balance of the relative abundance of tree species. Suppressing the more competitive vegetation, mainly Rubus fruticosus and graminoids, led to the recruitment and growth of tree species sensitive to competition and good tree species diversity. Practical recommendations in terms of vegetation control relative to tree regeneration and plant species richness are given.     

Effects of site preparation on emergence and early establishment of Larix olgensis in montane regions of northeastern China

Larix olgensis is one of the important commercial timber tree species in northeastern China, but it does not naturally regenerate in L. olgensis forests even after cutting. To identify the causes of the regeneration failure, the authors tested the effects of site preparation in a thinned stand and in a clearcut on the emergence and early establishment of L. olgensis seedlings. The results indicated that there was no correlation between cumulative emergence percentage (CEP) of L. olgensis seedlings and canopy openness, but the CEP was significantly affected by site preparation (P < 0.001). The average seedling survival rates after treatments consisting of removing both litter cover and understory vegetation cover were significantly higher in the clearcut than in the thinned stand (P < 0.001). Without site preparation, all seedlings in both the thinned stand and the clearcut disappeared within the first growing season, meaning that ground cover (understory vegetation cover and litter cover) was not beneficial to seedling survival. The average growth of seedlings exhibited significant differences between the thinned stand and the clearcut, and among the site preparations (P < 0.001). Intensely thinned stands without ground cover improved seedling growth. These results imply that light and ground cover might be the main factors limiting the early establishment of L. olgensis seedlings in managed plantation forests     

Spatio-temporal variation of environmental signals inducing seed germination in temperate conifer plantations and natural hardwood forests in northern Japan

To document the spatial and temporal variation of environmental signals inducing seed germination in temperate forests, we measured temporal patterns of environmental signals and seed germination of six pioneer tree species in unthinned and thinned stands of conifer forests (Cryptomeria japonica plantations) and in the understory and gaps of hardwood forests in Japan. We also conducted germination experiment in laboratory for the six pioneer species to test the effects of red:far-red (R:FR) light ratio and temperature fluctuations on the seed germination. In conifer forests, the photosynthetic photon flux density (PPFD), the R:FR ratio, and the amplitude of temperature fluctuations in thinned stands were 2, 1.5, and 3 times higher, respectively, than those of unthinned stands. The PPFD and R:FR ratios just above forest floor also increased after the removal of thick litter accumulation. As a result, higher seed germination was observed in thinned compared to unthinned stands for three photoblastic species, whereas little differences were observed for three non-photoblastic species. These findings suggest that thinning, which frequently reduces litter accumulation, can substantially affect the regeneration of pioneer species and the resultant species diversity in conifer plantations. None of the measured environmental signals changed seasonally in unthinned stands of conifer forests, but they all changed remarkably in the understory of the hardwood forests. In this system, all signals were high and nearly identical to those in the gaps in early spring prior to canopy closure. Thus, the percent germination of the three photoblastic species was enhanced by high R:FR ratios and/or large temperature fluctuations even beneath the canopy and was nearly equal to that in the thinned conifer stands where the environmental conditions were nearly identical to those in the gaps. However, all of the environmental signals decreased with the expansion of canopy leaves and reached minimums at canopy closure. Even in the thinned stands and the gaps, the PPFD and magnitude of temperature fluctuations decreased over time due to shading by growing herbs and/or emerging canopy leaves. In these temporally changing environments, the germination of all photoblastic species ceased simultaneously. This study clearly demonstrated that the environmental signals inducing seed germination of photoblastic pioneer species spatially and temporally change in temperate forests, particularly in deciduous hardwood forests. Furthermore, these signals, PPFD, R:FR ratio, and the amplitude of temperature fluctuations, appear to play a very important role in tree regeneration and subsequent species diversity.     

Do harrowing and fertilisation at middle rotation improve tree growth and site quality in <Emphasis Type="Italic">Eucalyptus globulus</Emphasis> Labill. plantations in Mediterranean conditions?

Effects of harrowing and fertilisation on tree growth, understory vegetation, forest floor litter and soil properties were studied in a field experiment installed in a 5-year-old first rotation eucalypt plantation. The treatments were harrowing (H), fertilisation (F), harrowing and fertilisation (HF) and a control without any treatments (C), with four replicates. Tree growth, root mass, tree nutrition status, understory vegetation, mass of forest floor litter and soil physical and chemical properties were monitored till the end of the rotation (stand was 14-year old). Effects of treatments on tree growth, root mass and timber production were not significant. Root nutrient concentration differed between treatments only few months after the beginning of the experiment, while differences regarding specific root length and area were observed during the first year. Harrowing (H and HF) led to significantly smaller biomass of understory vegetation (<13.4 g m⁻²) than other treatments (33−61 g m⁻²) during the first 3 years, but at the end of the experiment differences were less important. Forest floor litter mass after 3 years and at the end of the experiment were similar among treatments. Soil bulk density significantly decreased by harrowing (H and HF treatments), but 16 months after treatments were similar. Although soil chemical properties were not significantly affected by treatments, a decrease in base cations occurred during the rotation period. Fertiliser application is a questionable practice under the economic point of view, whereas harrowing can reduce the fuel load only during a short period, without negative effects on productivity and soil quality.     

Understory bamboo flowering provides a very narrow light window of opportunity for canopy-tree recruitment in a neotropical forest of Misiones, Argentina

Chusquea ramosissima is a native monocarpic bamboo species growing in subtropical forests of northeastern Argentina, which can dominate gaps and open forests in the region, particularly after human disturbance. This bamboo species started to flower in different areas of northeastern Argentina in 2001, with the flowering peak during 2002 and 2003 and small isolated flowering events still occurring until 2010. We studied the effects of C. ramosissima flowering and die-back on microclimate, litter decomposition, nutrient availability, sapling growth, abundance and regeneration of tree canopy species. We wanted to know how environmental conditions and ecosystem processes change through time after bamboo flowering and if bamboo die-back would favor regeneration of canopy trees. Twenty 50 × 50 m plots of flowering and non-flowering bamboo were permanently marked and vegetation dynamics as well as nutrient cycling and microclimate studies were performed. C. ramosissima die-back enhanced growth and reduced mortality rate of tree saplings during the first year after flowering. Only growth of tree saplings previously established was enhanced by the flowering event and tree-species richness and saplings abundance of canopy trees did not change as expected due to bamboo flowering. The short-term effect of tree saplings growth was likely due to incident solar radiation at the forest floor which doubled in the first year after the bamboo flowering event. Increased light availability at the forest floor simultaneously promoted the growth of other understory plants such as ferns, lianas and Piper spp. that rapidly colonized gaps and intercepted a percentage of the incident solar radiation after the first year, which together with an increased litter layer due to the senescence of the bamboo, may have inhibited establishment of new tree individuals and affected tree growth. Contrary to predictions, soil water, litter decomposition and soil nutrients were not significantly affected by bamboo flowering. Thus, successful tree regeneration in gaps following bamboo flowering appears to be restricted to a very narrow window of increased light availability (i.e., 1 year) before growth of other understory plants and rapid re-colonization of bamboo. Changes in resource availability, and the opportunity for overstory regeneration after bamboo flowering events appears to depend on climatic and community characteristics of the ecosystem where the flowering event occurs and also, on the flowering patterns and their synchronicity.     

Canopy tree development and undergrowth bamboo dynamics in old-growth Abies–Betula forests in southwestern China: a 12-year study

Interactions between forest canopy characteristics and plants in the forest understory are important determinants of forest community structure and dynamics. In the highlands of southwestern, China the dwarf bamboo Bashania fangiana Yi is an understory dominant beneath a mixed canopy of the evergreen Abies faxoniana (Rheder & Wilson) and the deciduous Betula utilis (D. Don). The goal of this study was to better understand the role of bamboo dominance, canopy characteristics, and periodic bamboo dieback on forest development. To achieve this goal, we measured tree seedling, tree saplings, and trees, forest canopy characteristics, and bamboo cover in permanent forest (n = 4) and gap plots (n = 31) in a mixed A. faxoniana and B. utilis forest in Sichuan, China. Dwarf bamboos died off in 1983 in the gap plots, and in three of the four forest plots. Forest development was assessed for the period 1984–1996. The seedling bank in forest and gap plots increased after bamboo die-off. A. faxoniana seedlings increased more than B. utilis in forest plots; the opposite pattern characterized gap plots. The proportion of seedlings on raised micro-sites on the forest floor also changed and new seedling were more abundant on the forest floor. By 1996, bamboo seedling cover and biomass had recovered to ca. 45% or their pre-flowering values. Rates of bamboo seedling recovery were faster beneath canopy gaps and deciduous trees than beneath forest or evergreen trees. Tree mortality exceeded recruitment in plots with dense bamboo; the opposite pattern was found in the plot with little bamboo. The mortality rate for B. utilis trees (2.4% year⁻¹) was higher than that for A. faxoniana (0.8% year⁻¹) and forests with dense bamboos became more open over the census period. Tree mortality was size-dependent and intermediate sized trees had the lowest rates of mortality. Stand basal area increased mainly due to greater basal area gain than loss for A. faxoniana. Interactions between tree species life history, canopy type, and bamboo life-cycles create heterogeneous conditions that influence tree and bamboo regeneration and contribute to the coexistence of A. faxoniana and B. utilis in old-growth forests in southwestern China.     

Role reversal in the stand dynamics of an angiosperm–conifer forest: Colonising angiosperms precede a shade-tolerant conifer in Afrotemperate forest

In mixed angiosperm–conifer forests worldwide, infrequent landscape-level catastrophic disturbances create a mosaic of persistent and different aged forest stands in the landscape with varying levels of dominance by the conifer component. In the ‘temporal stand replacement model’ (TSRM), disturbance creates conditions favouring a colonising cohort that is replaced by a suite of relatively shade-tolerant canopy species, which establish following the synchronous senescence of the pioneer canopy. In most southern hemisphere mixed angiosperm–conifer forests, with the exception of those in southern Africa, the establishing cohort is usually a large and very long-lived (550–650 years) conifer that is gradually replaced by angiosperms. As an explanation of the apparent dominance of the conifer Podocarpus latifolius, we examine the efficacy of the TSRM in mixed Afrotemperate forests where the establishing cohort is not a conifer. Forest succession in Afrotemperate forests was deterministic with the successive replacement of species determined first by their establishment success in shaded environments, and second, by their relative longevity. Several angiosperm species that were common canopy dominants established a pioneer cohort but were gradually replaced by P. latifolius, a shade-tolerant species. Continuous regeneration beneath the angiosperm canopy by P. latifolius eliminates synchronous canopy senescence, a key feature of the TSRM, as a mechanism driving the temporal replacement of canopy species. Senescing angiosperms created canopy gaps that were colonised by grasses and ferns, which suppressed canopy tree regeneration. In contrast, with continuous regeneration beneath the shaded canopy, P. latifolius gains a critical advantage over angiosperms at gap formation. Thus, in the absence of fairly large-scale natural disturbances, conifers come to dominate Afrotemperate forests. Commensurate with the latter, conifers in Podocarpus-forest were dated to approximately 320 years, more than 100 years older than the oldest P. latifolius in angiosperm-dominated forest. Tree life-history differences (shade tolerance, longevity) and the time since disturbance drive successional change from an angiosperm-dominated system to a stage dominated by P. latifolius. In general, the TSRM is a plausible explanation for the observed canopy tree structure and dynamics in mixed Afrotemperate forests. South African Afrotemperate forest is unusual among other southern hemisphere mixed angiosperm–conifer forests in that a suite of angiosperm canopy species, rather than a single conifer species, forms the colonising cohort.     

Effects of gaps on regeneration of woody plants: a meta-analysis

Forest gaps, openings in the canopy caused by death of one or more trees, have a profound effect on forest regeneration and drive the forest growth cycle. It is therefore necessary to understand the effects of forest gaps on regeneration for modern forest management. In order to provide a quantitative assessment of the effects of forest gaps on regen-eration of woody plants, we conducted this review of gap effects on woody plant regeneration on the basis of 527 observations from 42 indi-vidual papers, and reported the results of these data in a meta-analysis. Overall, densities of regenerated woody plants were significantly greater （359%） in forest gaps than on the closed-canopy forest floor. The regen-eration density in gaps of plantation forests was significantly greater （P＆lt;0.05） than that of natural forest because the regeneration in gaps of plan-tation forests was improved by both gap effects and experimental meas-ures. Similarly, in comparison to natural gaps, regeneration was better enhanced in artificial gaps. Regeneration density exhibited a significantly positive correlation with gap size, but a negative correlation with gap age because the gap size decreased with increasing gap age. Shade tolerance of woody plants affected regeneration density in gaps and understory. Average regeneration density of shade-tolerant species exhibited a sig-nificantly positive response to gaps but densities remained lower in total than those of intermediate and shade-intolerant species. Gap effects on regeneration decreased in response to increasing temperature and pre-cipitation because of the limiting effects of lower temperature and moisture on woody plant regeneration. In summary, forest gaps enhance woody plant regeneration, and the effects of gaps varied by forest type, gap characteristics, environmental factors and plant traits. The results of this meta-analysis are useful for better understanding the effects and roles of gaps on forest regeneration and forest management.     

Understory plant communities of boreal mixedwood forests in western Canada: Natural patterns and response to variable-retention harvesting

We examined patterns of variation in richness, diversity, and composition of understory vascular plant communities in mixedwood boreal forests of varying composition (broadleaf, mixedwood, conifer) in Alberta, Canada, before and for 2 years following variable-retention harvesting (clearcut, 20 and 75% dispersed green tree retention, control). Broadleaf-dominated forests differed from mixedwood or conifer-dominated forests in that they had greater canopy cover, litter depth, soil nitrogen, warmer soils, as well as greater shrub cover, herb and shrub richness and diversity (plot scale). In contrast, conifer, and to a lesser extent mixedwood, forest had greater β diversity than broadleaf forest. Overall, mixedwood and conifer forests were similar to one another, both differed from broadleaf forest. Several species were found to be significant indicators of broadleaf forest but most of these also occurred in the other forest types. Understory composition was related to canopy composition and edaphic conditions. Variable-retention harvesting had little effect on understory cover, richness, or diversity but resulted in reduced richness and β diversity at a larger scale. The clearcut and 20% treatments affected composition in all forest types. Early successional species and those common in disturbed sites were indicators of harvesting while evergreen, shade-tolerant understory herbs were indicators of the control forest and 75% retention harvest. We conclude that it is important to maintain a range of variation in canopy composition of mixedwood forests in order to conserve the associated understory communities. The presence of conifers in these forests has a particularly important influence on understory communities. The threshold for a lifeboat effect of variable-retention harvesting is between 20 and 75% retention. Examination of richness and β diversity at a variety of scales can provide interesting information on effects of harvesting on spatial reorganization and homogenization of understory plant communities.     

模拟林冠损伤对藜蒴栲群落林下植被生物多样性的影响

以2008年冰雪灾害对森林生态系统的破坏为背景,通过人工去除林冠层保留树干模拟森林的机械损伤,研究粤北小坑流域藜蒴栲群落林下植被的变化.结果表明:(1)试验1.5年后,去除林冠层、林下添加枯枝落叶的处理,林下灌木、草本科、种及乔木幼苗种的数量显著增加(P＜0.05);去除林冠层且地表枯落物层保持不变的处理,林下草本科、种数量增加显著,乔木幼苗及灌木科、种差异不显著(P＞0.05);林下添加枯枝落叶,林冠层不做处理,林下植被各生活型物种数有所减少,但差异不显著.(2)去除林冠层后,一些阳生种如野桐、山乌桕、山苍子、红紫珠、蕨状苔草、广东蛇葡萄等大量入侵并占据优势地位,林下植被盖度显著提高.(3)去除林冠层在短期内可显著增加林下植被生物多样性,添加枯落物对林下植被生物多样性影响不显著.