Succession stage variation in population size in an early-successional herb in a peri-urban forest

Urban and peri-urban forests incur high anthropogenic pressures (e.g. recreational activities, artificialization, and eutrophication). Plant species from early-successional, transient, forest habitats, often characterized by a short life span and a persistent seed bank in the soil may differ from late-successional species in key-factors for population persistence. This study investigated variation in population size and seedling recruitment for different forest succession stages and three consecutive years in Centaurium erythraea, an early-successional biennial herb, occurring in a peri-urban forest of Brussels urban zone (Belgium). Forest succession stage had a significant impact on C. erythraea population size and on its temporal fluctuation. Populations in closing vegetation (evolving to late-succession stages) showed small population sizes and a low number of recruits compared to populations from stable early-succession vegetation and clearcuts. The number of recruits was the highest after clearcutting, which can be related to the expression of the soil seed bank. Populations showed year-to-year variation in size (flowering individuals and recruits), even in stable (over three years) early-succession forest vegetation. In the absence of disturbance changing succession stage, population size is expected to depend on seed set of the previous years and subsequent seedling recruitment, which can be affected by environmental stochasticity. Opening gaps in the herbaceous vegetation may stimulate seedling recruitment, also in unoccupied patches where “cryptic” seed populations are present in the soil. Forest path and road verges, despite their potential negative impact on forests, can constitute refuge habitats for early-successional forest plant species. Their management should involve the preservation of these species.     

Recovery of late-seral vascular plants in a chronosequence of post-clearcut forest stands in coastal Nova Scotia,Canada

We investigated the impacts of clearcutting on the ground vegetation of remnant late-successional coastal Acadian forests in southwestern Nova Scotia. Vegetation was sampled in 750 1-m² quadrats established in 16 stands belonging to different recovery periods since clearcutting (3–54 years) and 9 late-successional forests (100–165 years) with no signs of significant human disturbance. Our objectives were to: i) describe the changes in species richness, diversity, and abundance of ground vegetation after clearcutting; ii) examine the responses of residual species (i.e., late-successional flora) to clearcutting; and iii) determine whether any forest species were restricted to or dependent upon the late-successional stages of stand development for maximal frequency and/or abundance. Although clearcutting had no immediate impact on overall alpha richness or diversity, the richness and diversity of residual plants declined after canopy removal and showed no evidence of recovery over 54 years of secondary succession. Consequently, compositional differences between secondary and late-seral stands persisted for many decades after clearcutting. Several understory herbs (e.g., Coptis trifolia (L.) , Oxalis montana (L.), Monotropa uniflora (L.)) were restricted to or attained their highest frequency and abundance in late-seral forests. These results suggest that the preservation of remnant old stands may be necessary for the maintenance of some residual plants in highly disturbed and fragmented forest landscapes in eastern Canada.     

Wind disturbance in remnant forest stands along the prairie-forest ecotone,Minnesota, USA

Strong winds are an important disturbance agent in northern Minnesota forests. On June 19, 1994, strong winds (>160 km h^(-1)) associated with a tornado damaged forested areas within the Rydell National Wildlife Refuge, situated in Polk County Minnesota along the prairie-forest boundary. Field sampling was conducted immediately following the storm to quantify the type and extent of damage in four different community types, and to project future composition based on the nature of the storm damage and current understory characteristics, including the impact of overbrowsing by deer. Basal area in six sampled remnant forest stands was reduced by 33.5%, although the damage was heterogeneous; basal area in one stand was reduced by 68.1%. The overall effect of the storm was the removal of early- successional species (primarily Populus tremuloides) in larger size classes. Trees situated at stand edges were not more susceptible to snapping or uprooting than interior trees. Projections of future stand composition indicate that wind disturbance, unlike other agents of disturbance such as fire, may accelerate succession on the Refuge, such that early-successional stands will assume a later-successional character, while Acer-Tilia stands should maintain their late-successional character. Overbrowsing and preferential foraging by deer may significantly alter stand recovery patterns.     

Effects of Previous Land-Use on Plant Species Composition and Diversity in Mediterranean Forests

At some point in their history, most forests in the Mediterranean Basin have been subjected to intensive management or converted to agriculture land. Knowing how forest plant communities recovered after the abandonment of forest-management or agricultural practices (including livestock grazing) provides a basis for investigating how previous land management have affected plant species diversity and composition in forest ecosystems. Our study investigated the consequences of historical “land management” practices on present-day Mediterranean forests by comparing species assemblages and the diversity of (i) all plant species and (ii) each ecological group defined by species’ habitat preferences and successional status (i.e., early-, mid-, and late-successional species). We compared forest stands that differed both in land-use history and in successional stage. In addition, we evaluated the value of those stands for biodiversity conservation. The study revealed significant compositional differentiation among stands that was due to among-stand variations in the diversity (namely, species richness and evenness) of early-, intermediate-, and late-successional species. Historical land management has led to an increase in compositional divergences among forest stands and the loss of late-successional forest species.     

Riverscape-level patterns of riparian plant diversity along a successional gradient, Willamette river, Oregon

Riparian forest communities dominated by Populus balsamifera ssp. trichocarpa L. (Torr. and Gray ex Hook.) Brayshaw are important contributors to biodiversity in terrestrial and aquatic ecosystems of the Western United States. Species composition along a successional gradient from stand initiation to late-succession of P. balsamifera-dominated riparian forests was investigated along 145 km of the Willamette River, Oregon. There were 151 total species encountered across 28 stands and a mean species richness of 33.3 species per stand. Young stands were dominated by P. balsamifera and Salix tree spp. and opportunistic herbaceous species. Understory trees, shrubs, and herbaceous species as well as late-successional tree species established 12–15 years after stand initiation. Fraxinus latifolia Benth. was the dominant late-successional tree species. Vertical structural diversity, P. balsamifera mean diameter at breast height, large tree biomass, and stand age were strongly correlated with understory species presence and abundance based on non-metric multidimensional scaling (NMS) ordination. There were no young stands on mid and high terraces and this was reflected in geomorphic position being strongly correlated with the stand age gradient. Abundance of Phalaris arundinacea L. an invasive grass species, was also significantly correlated with plant species composition and abundance. This study indicates that Willamette River riparian forests are diverse and therefore important to the biodiversity of the Willamette River valley and that their presence as a mosaic of communities of different successional stages may be threatened by human interventions, including influences exerted by introduced plant species.     

Bird communities in habitats along a successional gradient: Divergent patterns of species richness,specialization and threat

The long history of human influence on northern temperate landscapes has created a mosaic of successional stages, from closed forest to open grassland. Various species thus adapted to different habitats and it is interesting to explore how these differences in species composition among particular successional stages translate into differences at the community level. For this purpose, we surveyed breeding birds in 233 patches of five different habitats covering a gradient from bare ground to forest in 29 abandoned military training sites scattered throughout the Czech Republic. Linear mixed effects modelling revealed that late-successional habitats (dense scrubland and forest) were the most species-rich, whereas early-successional stages hosted bird communities with the highest habitat specialization and threat level. These results suggest that the habitats of late-successional stages are important for the maintenance of high bird species richness, but that early-successional habitats are essential for highly specialized and threatened bird species. Given the highly adverse impacts of agricultural intensification and land abandonment on open habitats, it is necessary to promote factors creating initial successional stages suitable for specialized and threatened species.     

Succession May Maintain High Leaf Area: Sapwood Ratios and Productivity in OldSubalpine Forests

Old forests are generally believed to exhibit low net primary productivity (NPP) and therefore to be insignificant carbon sinks. This relationship between age and NPP is based, in part, on the hypothesis that the biomass of respiratory tissues such as sapwood increases with age to a point where all photosynthate is required just to maintain existing tissue. However, this theoretical connection between respiration:assimilation ratios and forest productivity is based on age-dependent trends in the sapwood:leaf ratios of individual trees and even-aged stands; it does not take into account such processes in natural forests as disproportional increases in shade-tolerant species over time and multiple-age cohorts. Ignoring succession and structural complexity may lead to large underestimates of the productivity of old forests and inaccurate estimates of the ages at which forest productivity declines. To address this problem, we compared biomass allocation and productivity between whitebark pine, a shade-intolerant, early-successional tree species, and subalpine fir, a shade-tolerant, late-successional species, by harvesting 14 whitebark pines and nine subalpine firs that varied widely in dbh and calculating regression models for dbh vs annual productivity and biomass allocation to leaves, sapwood, and heartwood. Late-successional subalpine fir allocated almost twice as much biomass to leaves as early-successional whitebark pine. Subalpine firs also had a much lower allocation to sapwood and higher growth rates across all tree sizes. We then modeled biomass allocation and productivity for 12 natural stands in western Montana that were dominated by subalpine fir and whitebark pine varying in age from 67 to 458 years by applying the regressions to all trees in each stand. Whole-stand sapwood:leaf ratios and stand productivity increased asymptotically with age. Sapwood:leaf ratios and productivity of whitebark pine in these stands increased for approximately 200–300 years and then decreased slowly over the next 200 years. In contrast, sapwood:leaf ratios of all sizes of subalpine fir were lower than those of pine and productivity was higher. As stands shifted in dominance from pine to fir with age, subalpine fir appeared to maintain gradually increasing rates of whole-forest productivity until stands were approximately 400 years old. These results suggest that forests such as these may continue to sequester carbon for centuries. If shade-tolerant species that predominate late in succession maintain high assimilation-to-respiration ratios in other forests, we may be underestimating production in old forests, and current models may underestimate the importance of mature forests as carbon sinks for atmospheric CO₂ in the global carbon cycle. Received 16 February 1999; accepted 24 November 1999.     

Temporal variation in plant-soil feedback controls succession

Soil abiotic and biotic factors play key roles in plant community dynamics. However, little is known about how soil biota influence vegetation changes over time. Here, we show that the effects of soil organisms may depend on both the successional development of ecosystems and on the successional position of the plants involved. In model systems of plants and soils from different successional stages, we observed negative plant–soil feedback for early-successional plant species, neutral feedback for mid-successional species, and positive feedback for late-successional species. The negative feedback of early-successional plants was independent of soil origin, while late-successional plants performed best in late- and worst in early-successional soil. Increased performance of the subordinate, late-successional plants resulted in enhanced plant community diversity. Observed feedback effects were more related to soil biota than to abiotic conditions. Our results show that temporal variations in plant–soil interactions profoundly contribute to plant community assemblage and ecosystem development.     

Contrasting effects of habitat quantity and quality on moth communities in fragmented landscapes

Habitat loss is commonly identified as a major threat to the loss of global biodiversity. In this study, we expand on our previous work by addressing the question of how lepidopteran species richness and composition vary among remnants of North American eastern deciduous forest located within agricultural or pastoral landscapes. Specifically, we tested the relative roles of habitat quantity (measured as stand area and percent forest in the greater landscape) and habitat quality (measured as tree species diversity) as determinants of moth species richness. We sampled >19 000 individuals comprising 493 moth species from 21 forest sites in two forested ecoregions. In the unglaciated Western Allegheny Plateau, the species richness of moths with woody host plants diminished as forest stand size and percent forest in the landscape decreased, but the total species richness and abundance of moths were unaffected by stand size, percent forest in the landscape, or tree species diversity. In contrast, the overall species richness and abundance of moths in the glaciated North Central Tillplain were affected primarily by tree species diversity and secondarily by forest size. Higher tree species diversity may reduce species loss from smaller forest stands, suggesting that small, diverse forests can support comparable numbers of species to those in less diverse, large stands. Smaller forests, however, contained a disproportionate number of moth species that possess larvae known to feed on herbaceous vegetation. Thus, although woody plant feeding moths are lost from forests with changes in stand area, new species appear capable of recolonizing smaller fragments from the surrounding habitat matrix. Our study further suggests that when species replacement occurs, local patch size and habitat quality may be more important than landscape context in determining the community structure of forest Lepidoptera.     

黑龙江省大兴安岭林区火烧迹地森林更新及其影响因子

林火干扰是大兴安岭森林更新的影响因子之一,研究火烧迹地森林更新的影响因子(立地条件、火前植被、火干扰特征)对理解生态系统的结构、功能和火后演替轨迹具有重要意义。选取呼中及新林林业局55个代表性火烧样地,利用增强回归树分析法分析了火烧迹地森林更新的影响因素。结果表明:(1)立地条件是影响针、阔叶树更新苗密度的主要因素;海拔对针叶树更新苗密度的影响最大;坡度对阔叶树更新苗密度影响最大;(2)距上次火烧时间对针叶树更新苗比重影响最大,其次是林型;(3)中度林火干扰后森林更新状况好于轻度和重度火烧迹地。根据火烧迹地森林更新调查分析可知:林型影响火后演替模式,火前为针叶树或阔叶树纯林,火后易发生自我更新(火后树种更新组成与火前林型相同),而针阔混交林在火干扰影响下易于发生序列演替(火后初期以早期演替树种更新为主)。     

Changes in plant species diversity along a chronosequence of vegetation restoration in the humid evergreen broad-leaved forest in the Rainy Zone of West China

Plant species diversity has been recognized as one of the vital attributes for assessing vegetation restoration. Changes in the diversity may be related to different stages of succession. In this study, 54 sites of humid, evergreen, broad-leaved forest were selected in the Rainy Zone of West China. A chronosequence of the sites was used to study the successive patterns of the diversity in the forest that had undergone natural regeneration for 5 to 350 years and to test the hypothesis that the diversity is maximized in mid-succession. Data were collected simultaneously at different stages of succession, and four α-diversity indices (species richness, Margalef index, Shannon-Wiener index, Pielou Evenness index) and two β-diversity indices (Whittaker index, Sørensen’s index) were calculated for each stratum in each plot. A total of 394 vascular plant species were recorded. From the β-diversity indices, the forest succession may be divided into the early-successional stage (before 50 years), mid-successional stage (from 50 to 300 years), and late-successional stage (after 300 years). In this community, the species diversity and richness were found to be the greatest at the mid-successional stage, followed by the late- and early-successional stages. The results of regression analysis indicated that the richness and Margalef index peaked around the 175th and 165th year, respectively. Shannon-Wiener index values also appeared to follow an approximately humped pattern of succession and were maximal around the 100th year. However, the species evenness did not show any significant relationship with successional age. Our results demonstrate (1) forest restoration is a long-term process and the formation of climax forest requires at least 300 years and (2) the forest has a strong capacity for restoration. Our results also suggest Lindera limprichitii and Machilus pingii as ideal tree species for afforestation because of their wide niche.     

Community composition and tree population structure in a sub-tropical broad-leaved forest along a disturbance gradient

Floristic composition, diversity, dominance and distribution pattern of species and tree population structure were studied in three stands of a sub-tropical wet hill broad-leaved forest of Meghalaya, India, along a disturbance gradient. Tree species diversity declined with increasing disturbance. Disturbed stands showed low equitability or high dominance and the undisturbed stand exhibited high equitability or low dominance. Contagious distribution among the tree species increased with increasing intensity of disturbance. Species showing regular distribution were restricted only to the undisturbed stand. Effect of disturbance on tree population structure was analysed using density-diameter curves. In the disturbed stands tree species showed reverse J-shaped and/or negative exponential curves, while those in the undisturbed stand exhibited sigmoid to bimodal mound shaped curves.     

Seed-bank convergence under different tree species during forest development

Seed banks are of vital importance for local plant persistence and recruitment, for maintaining both plant and genetic diversity and for habitat restoration. Yet, seed-bank dynamics, particularly on the long term and in deciduous forests, remain poorly understood. Additionally, information on compositional seed-bank differences under contrasting tree canopies remains scarce.This study aims at quantifying long-term seed-bank dynamics by sampling vegetation and seed banks along a four-stage successional chronosequence (40, 80, 120 and 250 years) using 12 10 m×10 m plots per forest stand age–class under fully developed oak–hornbeam and beech canopies.Seed banks were remarkably abundant and diverse. Species richness and seed density declined steeply with forest stand age, regardless of canopy species. Seed-bank composition differed significantly with stand age, yet also with tree species. Most likely, tree species-dependent ecosystem engineer effects on light availability and possibly also litter quality affect the seed bank through the vegetation. Compositional differences between seed banks from stands with a different canopy diminished with increasing stand age, possibly due to a gradual loss of species with a less persistent seed bank.Long-term seed-bank dynamics in deciduous forests seem to consist predominantly of a unidirectional and predictable depletion of the seed bank as long as large disturbances, which would allow seed-bank replenishment of early-successional species, are lacking. Furthermore, forest seed-banks appear to converge upon a characteristic seed bank in the later stages of forest development, irrespective of canopy composition, driven by seed-bank depletion and limited input from the herb layer.     

Variation in protein complexation capacity among and within six plant species across a boreal forest chronosequence

We investigated among and within species variation in several litter chemical properties, including protein complexation capacity (PCC), for six plant species across a boreal forest chronosequence in northern Sweden across which stand fertility declines sharply with stand age. We hypothesized (1) that evergreen species which dominate in late-successional stands would exhibit higher PCCs than deciduous species that dominate in young stands, (2) that individual species would increase their PCCs in response to nutrient limitation as succession proceeds, and (3) that differences in PCC among litter types would determine their interactive effects with proteins on soil N and C mineralization. The data demonstrated a high PCC, but a low PCC per unit of soluble phenol, for two deciduous species that dominate in early-successional high fertility stands, providing mixed support for our first hypothesis. No species demonstrated a significant correlation between their PCC and stand age, which did not support our second hypothesis. Finally, a soil incubation assay revealed that litter extracts for three of the six species had negative interactive effects with added proteins on N mineralization rates, and that all six species demonstrated positive interactive effects with protein on C mineralization. This pattern did not provide strong support for our third hypothesis, and suggests that N immobilization was likely a more important factor regulating N mineralization than stabilization of proteins into tannin complexes. These data suggest that multiple interactive mechanisms between litter extracts and proteins likely occur simultaneously to influence the availability of N in soils.     

宝天曼地区栓皮栎林恢复过程中高等植物物种多样性变化

对宝天曼地区不同恢复阶段栓皮栎林的高等植物物种多样性特征进行了初步分析。结果表明,各层物种丰富度和多样性指数在不同恢复阶段栓皮栎林表现出草本层＞灌木层＞乔木层的趋势。乔木层物种丰富度和多样性指数基本上随恢复时间的增加而增加;灌木层在未破坏的栓皮栎林内较低,而在其他恢复阶段的栓皮栎林内基本相同;草本层在恢复５年、１５年和２５年的栓皮栎林中较高,而在恢复４５年和未破坏的栓皮栎林内较低。乔木层物种均匀度指数在未破坏的栓皮栎林中较高,在恢复２５年的栓皮栎林中较低;灌木层在恢复４５年的栓皮栎林中较高,在恢复１５年的栓皮栎林较低;草本层在恢复５年的栓皮栎林中较高,在恢复１５年的栓皮栎林中较低。不同恢复阶段栓皮栎林各层次间的物种多样性差异大多不显著,只有乔木层和草本层及灌木层和草本层之间的物种丰富度指数有显著差异。不同恢复阶段栓皮栎林在乔木层物种多样性特征上的差异最大,在灌木层物种多样性特征上有一定差异,在草本层物种多样性特征上没有明显的差异。     

Effects of canopy gaps on the growth of tree seedlings from subtropical broad-leaved evergreen forests of southern China

Seedling performance, morphology, and leaf characteristics of evergreen tree species from different successional stages of Chinese broadleaved evergreen forests were studied in four simulated gap environments: in 100, 55, 33, and 18% of full sunlight. The first two represent the light regimes of clearings (gaps secondarily become more open due to human activity and natural erosion). The hypothesis was tested that early-successional species achieve their greatest seedling size in clearings and contrast in this respect with late-successional species. Late-successional species were expected to show a stronger morphological response (leaf/root ratio) to clearings than early-successional ones. The results provided some evidence in support of these hypotheses with one exception, namely that late-successional Castanopsis fargesii appeared to be a highly sun-tolerant species. It is suggested that Castanopsis seedlings are competitive in large gaps and clearings.     

Differences in bird communities in postfire silvicultural practices stands within pine forest of South Korea

We examined differences in bird communities in relation to characteristics of habitat structure in a pine forest, Samcheok, South Korea. An unburned stand, a stand burned 7 years earlier and then naturally restored, and a stand where Japanese red pine Pinus densiflora seedlings were planted after the fire were used for the survey. Habitat structure was dramatically changed by postfire silvicultural practices. Number of stand trees, shrubs, seedlings, snags, and vegetation coverage were significantly different among study stands. We made 1,421 detections of 46 bird species during 23 separate line transect surveys per stand between February 2007 and December 2008. The mean number of observed bird species and individuals, bird species diversity index (H′), and Simpson’s diversity index (D _s) were highest in the unburned stand and lowest in the pine seedling stand. There were more species and individuals of forest-dwelling birds in the unburned stand than both burned stands. Canopy and cavity nesters, foliage searchers, bark gleaners, and timber drillers were significantly higher in the unburned stand. In the pine seedling stand, densities of birds that prefer open field and shrub cover were higher. Stand structure was simplified in the pine seedling stand by postfire practices. Because of differences in habitat structure and bird communities, postfire practices in the burned stand should be re-evaluated. Also, management strategies for pine forest after forest fires are needed based on results of long-term experiments.     

Stand and neighbourhood parameters as determinants of plant species richness in a managed forest

Abstract. Patterns of species richness in a managed forest were related to ecological factors at two spatial scales. Local variables within a plot (5 m × 5 m) included 13 soil classes and nine stand classes. Neighbourhood variables were defined within a 25 m radius from the focal plot and were measured using a Neighbourhood Diversity Index (NDI - total diversity of different stand classes) and nine Neighbourhood Scores (NS - relative dominance of a given stand class). 224 species were registered in the survey of which 138 were true forest species. Local parameters (stand and soil class) were weak predictors of total and forest species richness. High total and forest species richness were significantly related to high values of NDI (i.e. heterogeneous neighbourhoods). Regression trees showed that total species richness was best predicted by the amount of roadside habitat in the neighbourhood and NDI. Forest species richness was positively related to NDI and negatively related to the amount of coniferous stands in the neighbourhood.     

Floristic diversity and co-occurrences in a subtropical broad-leaved forest and two contrasting regrowth stands in central-west Yunnan Province,China

Many of the natural forested ecosystems that still remain in mainland China are being cleared with potentially detrimental effects on woody plant species diversity on both local and regional scales. The most extensive stand of subtropical broad-leaved forest remaining in China is located in Yunnan Province. In an effort to document the influence of human-induced disturbance on Yunnan's woody flora, floristic inventories were conducted in a stand of primary forest and in regrowth stands located in its interior and along its outer margin in the Xujiaba Nature Sanctuary in the Ailao Mountain Range. Of particular interest was the location of the disturbance relative to the primary forest source area. A total of 134 woody plant species representing 74 genera and 43 families were recorded. The floristics of the two regrowth stands were significantly different from each other, with < 10% of their respective floras comprised of co-occurring species. The interior regrowth stand had a higher number of co-occurring species with the primary forest; however, > 40% were still non-co-occurring.The principal families represented in the primary forest and the interior regrowth stand were Aquifoliaceae, Berberidaceae, Fagaceae, Lauraceae, Rosaceae, Smilacaceae, Symplocaceae, Theaceae, and Vacciniaceae. The three dominant species with relative importance values ranging from > 5% to 18% in both the primary forest and the interior regrowth stand were Castanopsis wattii, Lithocarpus jingdongensis, and Symplocos sumuntia. The edge regrowth stands had the lowest species diversity and were dominated by the native pine Pinus yunnanensis, with a relative importance of 24%. The principal families represented in the edge regrowth stand were Betulaceae, Ericaceae, Fagaceae, Myricaceae, Pinaceae, and Theaceae. Only the Fagaceae and Theaceae were well-represented in all three stands. The results of the study document the low species diversity in post-cutting regrowth on the margins of the primary forest as compared with post-cutting regrowth in the forest interior.