Orthoptera as ecological indicators for succession in steppe grassland

Understanding the effects of land-use on threatened ecosystems is of special relevance for nature conservation. The aim of our study was to use Orthoptera as ecological indicators for succession in Central European steppe grasslands.Orthoptera showed a clear response to succession. Each successional stage harboured a unique assemblage. Species richness of habitat specialists was highest in the earliest seral stages. In contrast, density of all species peaked at the intermediate successional stage. Early successional stages are mostly likely to be preferred by specialized Orthoptera because they provide warm suitable oviposition sites (bare ground) and microclimatic conditions. The density peak in the mid-successional stage probably reflects a trade-off between favourable ambient temperatures for optimal development, sufficient food and shelter against predators.Although all successional stages of steppe grassland are relevant for conservation, early and mid-successional stages are the most important. Consequently, conservation management should aim at re-introduction of a traditional, low-intensive land use for abandoned steppe grasslands. As an optimal land use, we recommend traditional rough grazing with sheep and goats, which creates a heterogeneous habitat structure with bare ground, and avoids the accumulation of litter, favouring Orthoptera.     

Herbivory on Handroanthus ochraceus (Bignoniaceae) along a successional gradient in a tropical dry forest

This study determined the temporal patterns of herbivory on Handroanthus ochraceus (Cham.) Mattos (Bignoniaceae) along a successional gradient in a seasonally dry tropical forest (SDTF) in southeastern Brazil. We assessed the diversity of free-feeding herbivore insects (sap-suckers and leaf-chewers), leaf herbivory rates, leaf nitrogen content, phenolic compounds, and spider abundance through the rainy season in three different successional stages (early, intermediate, and late). Sampling was conducted in December, at the beginning of the rainy season (with fully expanded young leaves), February (mid-aged leaves), and April, at the end of rainy season (old leaves). Fifteen reproductive trees of H. ochraceus were sampled per successional stage in each month of sampling. Herbivore diversity was highest in the early stage of succession, but herbivory rates were highest in the intermediate and late stages. This result was probably related to differences in herbivore community composition and leaf quality across successional stages. The highest herbivore abundance was found in April in the early successional stage. In addition, we found low levels of herbivory in the intermediate and late successional stages in the second half of the rainy season. For each successional stage, leaf nitrogen content decreased through the rainy season, whereas the concentration of phenolic compounds increased. For the intermediate and late successional stages, the temporal changes that took place as the rainy season progressed corroborated the following hypotheses postulated for SDTFs: (1) both the abundance of chewing insects and herbivory rates decreased, (2) the abundance of natural enemies (i.e., spiders) increased, and (3) leaf quality decreased. These results suggest that the described herbivory patterns are robust for advanced successional stages (intermediate and late) of the SDTFs, but may not apply to early successional stages of these forests.     

松材线虫侵袭引发的植被演替对鸟类群落的影响

松材线虫(Bursaphelenchusxylophilus)是引起严重森林病害的外来物种,可能导致森林生态系统急剧退化。为了探讨在松材线虫侵袭所引起的森林演替过程中植被结构变化对鸟类群落的影响,我们研究了这些常绿阔叶演替林中的鸟类群落结构。2004年5–6月,我们在浙江省宁波市选择了分别处于3个不同演替阶段的7个样地:包括象山县被松材线虫侵袭5年后的常绿阔叶灌丛和侵袭12年后的常绿阔叶林样地各3个,同时在天童国家级森林公园选择了约40龄的常绿阔叶林样地1个。结果显示,在常绿阔叶灌丛和松材线虫侵袭12年后的常绿阔叶林中,鸟类物种多样性和丰富度均显著高于约40龄常绿阔叶林;3类栖息地的鸟类多度、均匀度均无显著差异;在侵袭12年后的常绿阔叶林中鸟类群落最丰富。多元回归分析结果显示,立地年龄是影响鸟类群落组成的主要因素。     

Plant diversity patterns in subtropical evergreen broad-leaved forests of Yunnan and Taiwan

The subtropical evergreen broad-leaved forests of Yunnan and Taiwan were compared along environmental and successional gradients with the aim of identifying important taxon and species diversity as well as the drivers of mountain biodiversity patterns. A detrended correspondence analysis of an exhaustive set of data collected from 105 and 223 plots for Yunnan and Taiwan, respectively, was applied to classify natural mature forest types. Additional data from 72 and 68 plots for Yunnan and Taiwan, respectively, were used for analyses of secondary succession. The floristic richness and diversity index were calculated for each type of forest. In Yunnan, the monsoon forests in mesic-humid sites had more taxa and tended to show higher species diversity than the other two forest types. In Taiwan, species diversity values were significantly higher in the Machilus–Castanopsis zone in the middle altitudes (500–1500 m) than for the other three forest zones. For both Yunnan and Taiwan, the forests at the middle successional stage showed significantly higher species diversity than those at the early successional stage. Differences in diversity between the middle and late stages were not significant. These findings highlight the high species diversity of the natural mature evergreen broad-leaved forests of both Yunnan and Taiwan. In the secondary forests, as succession proceeds, species diversity comes to resemble that of the natural mature forests. In both ecosystems, the drivers of species diversity patterns are moisture, altitude, and succession/disturbance.     

Does tree diversity increase wood production in pine forests?

Recent experimental advances on the positive effect of species richness on ecosystem productivity highlight the need to explore this relationship in communities other than grasslands and using non-synthetic experiments. We investigated whether wood production in forests dominated by Aleppo pine (Pinus halepensis) and Pyrenean Scots pine (Pinus sylvestris) differed between monospecific and mixed forests (2-5 species) using the Ecological and Forest Inventory of Catalonia (IEFC) database which contains biotic and environmental characteristics for 10,644 field plots distributed within a 31,944 km(2) area in Catalonia (NE Spain). We found that in Pyrenean Scots pine forests wood production was not significantly different between monospecific and mixed plots. In contrast, in Aleppo pine forests wood production was greater in mixed plots than in monospecific plots. However, when climate, bedrock types, radiation and successional stage per plot were included in the analysis, species richness was no longer a significant factor. Aleppo pine forests had the highest productivity in plots located in humid climates and on marls and sandstone bedrocks. Climate did not influence wood production in Pyrenean Scots pine forests, but it was highest on sandstone and consolidated alluvial materials. For both pine forests wood production was negatively correlated with successional stage. Radiation did not influence wood production. Our analysis emphasizes the influence of macroenvironmental factors and temporal variation on tree productivity at the regional scale. Well-conducted forest surveys are an excellent source of data to test for the association between diversity and productivity driven by large-scale environmental factors.     

The response of mammals to forest fire and timber harvest in the North American boreal forest

1. This paper reviews and compares the effects of forest fire and timber harvest on mammalian abundance and diversity, throughout successional time in the boreal forest of North America. 2. Temporal trends in mammal abundance and diversity are generally similar for both harvested and burned stands, with some differences occurring in the initiation stage (0–10 years post disturbance). 3. Small mammals and ungulates are most abundant immediately post disturbance, and decrease as stands age. Lynxes and hares utilize mid-successional stands, but are rare in young and old stands. Bats, arboreal sciurids and mustelids increase in abundance with stand age, and are most abundant in old growth. 4. Substantial gaps in the data exist for carnivores; the response of these species to fire and harvest requires research, as predator–prey interactions can affect mammal community structure in both early and late successional stages. 5. The lack of explicit treatment of in-stand forest structure post disturbance, in the reviewed literature made comparisons difficult. Where forest structure was considered, the presence of downed woody material, live residual trees and standing dead wood were shown to facilitate convergence of mammal communities to a pre-disturbance state for both disturbance types. 6. Mammalian assemblages differed considerably between successional stages, emphasizing the importance of maintaining stands of each successional stage on the landscape when implementing forest management strategies.     

Impact of a hurricane on the herpetofaunal assemblages of a successional chronosequence in a tropical dry forest

Land‐use change is the main cause of deforestation and degradation of tropical forest in Mexico. Frequently, these lands are abandoned leading to a mosaic of natural vegetation in secondary succession. Further degradation of the natural vegetation in these lands could be exacerbated by stochastic catastrophic events such as hurricanes. Information on the impact of human disturbance parallel to natural disturbance has not yet been evaluated for faunal assemblages in tropical dry forests. To evaluate the response of herpetofaunal assemblages to the interaction of human and natural disturbances, we used information of pre‐ and post‐hurricane herpetofaunal assemblages inhabiting different successional stages (pasture, early forest, young forest, intermediate forest, and old growth forest) of dry forest. Herpetofaunal assemblages were surveyed in all successional stages two years before and two years after the hurricane Jova that hit the Pacific Coast of Mexico on October 2011. We registered 4093 individuals of 61 species. Overall, there were only slight effects of successional stage, hurricane Jova or the interaction between them on abundance, observed species richness and diversity of the herpetofauna. However, we found marked changes in estimated richness and composition of frogs, lizards, and snakes among successional stages in response to hurricane Jova. Modifications in vegetation structure as result of hurricane pass promoted particular changes in each successional stage and taxonomic group (anurans, lizards, and snakes). Secondary forests at different stages of succession may attenuate the negative effects of an intense, short‐duration, and low‐frequency natural disturbance such as hurricane Jova on successional herpetofaunal trajectories and species turnover.     

Early succession of butterfly and plant communities on set-aside fields

 Hypotheses on secondary succession of butterfly and plant communities were tested using naturally developed 1- to 4-year-old set-aside fields (n = 16), sown fields (n = 8) and old meadows (n = 4) in 1992 in South Germany. Pioneer successional fields (1st and 2nd year of succession, dominated by annuals) and early successional fields (3rd and 4th year of succession where perennials, especially grasses became dominant) had fewer plant species than mid-successional fields (old meadows). In contrast to established hypotheses, mean number of plant species decreased from 1- to 4-year-old set-aside fields. Species richness of butterfly communities did not change during the first four years of succession, but species composition changed greatly. Pioneer successional fields were characterized by (1) specialized butterflies depending on annual pioneer foodplants (e.g. Issoria lathonia), and (2) species preferring the pioneer successions despite their host plants being more abundant on early and mid-successional fields (e.g. Papilio machaon). The variability in butterfly species richness was best explained by flower abundance which was closely correlated with plant species richness. Species whose abundance was correlated with habitat connectivity were significantly smaller than species which correlated with flower abundance. Numbers of caterpillar species were correlated with numbers of adult butterfly species. Life-history features of butterflies changed significantly from pioneer to early and mid-successional fields. We found decreasing body size and migrational ability, decreasing numbers of species hibernating as imago, decreasing numbers of generations and increasing larval stage duration with age of succession, but, contrary to expectation, host plant specialization, numbers of egg-cluster laying species and egg diameter did not change with successional age. Received 18 September 1995 / Accepted: 17 July 1996     

Distribution of Myxomycetes on coarse woody debris of <Emphasis Type="Italic">Pinus densiflora</Emphasis> at different decay stages in secondary forests of western Japan

The relationship between myxomycete species and the decay stage of wood of Pinus densiflora coarse woody debris was investigated in warm temperate secondary forests of western Japan. The number of species and species diversity of the myxomycete community reached the maximum on moderately decayed wood. The 25 dominant species recorded from 8 or more samples of the total 1530 samples were arranged in order of the succession index corresponding to the stage of decay. Species on slightly decayed hard pine wood were characterized by Stemonitis splendens, Enerthenema papillatum, and Physarum viride, whereas species of Cribrariaceae were found on brittle decayed soft wood increasing abundance according to the decay stages. Most of the species occurred where there was sufficient moisture preserved in the environment of the decaying wood, although S. splendens specifically emerged in low-moisture environments. Because the myxomycete species had preference to different decay stages of wood, it appears that they change sequentially during myxomycete community succession on dead pine wood according to the progression of decay.     

Succession Influences Wild Bees in a Temperate Forest Landscape: The Value of Early Successional Stages in Naturally Regenerated and Planted Forests

In many temperate terrestrial forest ecosystems, both natural human disturbances drive the reestablishment of forests. Succession in plant communities, in addition to reforestation following the creation of open sites through harvesting or natural disturbances, can affect forest faunal assemblages. Wild bees perform an important ecosystem function in human-altered and natural or seminatural ecosystems, as they are essential pollinators for both crops and wild flowering plants. To maintain high abundance and species richness for pollination services, it is important to conserve and create seminatural and natural land cover with optimal successional stages for wild bees. We examined the effects of forest succession on wild bees. In particular, we evaluated the importance of early successional stages for bees, which has been suspected but not previously demonstrated. A range of successional stages, between 1 and 178 years old, were examined in naturally regenerated and planted forests. In total 4465 wild bee individuals, representing 113 species, were captured. Results for total bees, solitary bees, and cleptoparasitic bees in both naturally regenerated and planted conifer forests indicated a higher abundance and species richness in the early successional stages. However, higher abundance and species richness of social bees in naturally regenerated forest were observed as the successional stages progressed, whereas the abundance of social bees in conifer planted forest showed a concave-shaped relationship when plotted. The results suggest that early successional stages of both naturally regenerated and conifer planted forest maintain a high abundance and species richness of solitary bees and their cleptoparasitic bees, although social bees respond differently in the early successional stages. This may imply that, in some cases, active forest stand management policies, such as the clear-cutting of planted forests for timber production, would create early successional habitats, leading to significant positive effects for bees in general.     

Is patience a virtue? Succession, light, and the death of invasive glossy buckthorn (Frangula alnus)

Non-native, invasive glossy buckthorn (Frangula alnus P. Mill.) threatens North American forests by inhibiting tree regeneration. While glossy buckthorn commonly invades younger, developing forests, we hypothesized that this species is competitively excluded as secondary succession proceeds. Specifically, we tested the hypothesis that glossy buckthorn mortality in New Hampshire forests is associated with low levels of photosynthetically active radiation (PAR), low levels of nutrients in the soil, and greater abundance of shade tolerant tree species. Twenty-six living and 26 dead glossy buckthorn individuals were randomly selected on three, mid-successional study sites in southeastern New Hampshire, USA. Shrub ages, indicated by wood ring counts, showed that both living and dead shrubs were members of the same cohort, roughly 16–18 years of age. Living and dead shrubs were compared as to (1) tree basal area in the surrounding forest (estimated by plot sampling), (2) light intensity in the vicinity of the buckthorn crowns (measured by Sunfleck Ceptometer), (3) canopy openness and % total transmitted PAR (estimated by hemispheric photography), and (4) concentrations of available Ca, P, Mg, and K in the B horizon. Relative to live buckthorn shrubs, dead individuals were associated with greater basal area of shade tolerant species, decreased PAR in the vicinity of shrub crown, and decreased soil concentrations of Ca, P, and Mg. The results are consistent with the hypothesis that competition for light and perhaps nutrients limits the ability of glossy buckthorn to persist in late successional stands. If these relationships are causal, active removal of glossy buckthorn might not be required to reduce its abundance in late successional stands.     

Changes in Forest Soil Properties in Different Successional Stages in Lower Tropical China

Background

Natural forest succession often affects soil physical and chemical properties. Selected physical and chemical soil properties were studied in an old-growth forest across a forest successional series in Dinghushan Nature Reserve, Southern China.

Methodology/Principal Findings

The aim was to assess the effects of forest succession change on soil properties. Soil samples (0–20 cm depth) were collected from three forest types at different succession stages, namely pine (Pinus massoniana) forest (PMF), mixed pine and broadleaf forest (PBMF) and monsoon evergreen broadleaf forest (MEBF), representing early, middle and advanced successional stages respectively. The soil samples were analyzed for soil water storage (SWS), soil organic matter (SOM), soil microbial biomass carbon (SMBC), pH, NH₄ ⁺-N, available potassium (K), available phosphorus (P) and microelements (available copper (Cu), available zinc (Zn), available iron (Fe) and available boron (B)) between 1999 and 2009. The results showed that SWS, SOM, SMBC, Cu, Zn, Fe and B concentrations were higher in the advanced successional stage (MEBF stage). Conversely, P and pH were lower in the MEBF but higher in the PMF (early successional stage). pH, NH₄ ⁺-N, P and K declined while SOM, Zn, Cu, Fe and B increased with increasing forest age. Soil pH was lower than 4.5 in the three forest types, indicating that the surface soil was acidic, a stable trend in Dinghushan.

Conclusion/Significance

These findings demonstrated significant impacts of natural succession in an old-growth forest on the surface soil nutrient properties and organic matter. Changes in soil properties along the forest succession gradient may be a useful index for evaluating the successional stages of the subtropical forests. We caution that our inferences are drawn from a pseudo-replicated chronosequence, as true replicates were difficult to find. Further studies are needed to draw rigorous conclusions regarding on nutrient dynamics in different successional stages of forest.     

Diversity and community composition of herbaceous plants in different habitat types in south‐east Cameroon

The composition of herbaceous vegetation was evaluated with the aim of characterizing forests at various ages of stand development. Herb stems were sampled in 250 4‐m² square plots distributed within six habitat types. A total of 36 herb species belonging to 15 families were recorded. Species richness did not significantly differ between habitat types. Most herb species occurred in all habitat types and were therefore generalists. However, a few indicator herb species were detected, and the results roughly suggested that herb species of the families Poaceae and Araceae were indicative of late successional forests; Zingiberaceae are indicative of early successional forests; and Commelinaceae, Costaceae, Cyperaceae and Marantaceae are indicators of flooded habitats. Species diversity and stem density of herbaceous plants did not change with forest succession as a decrease in abundance and frequency of occurrence of pioneer species in late successional forests was counterbalanced by the presence of generalist and late successional species. However, increasing proportions of dwarf stems in late successional forests translated to changes in the vertical structure of herbaceous plant communities. Herbivory pressure by gorillas did not have a notable effect on herbaceous plant community development. This study contributes to the definition of herbaceous ecological indicators of forest succession in different settings.     

Succession of secondary forests affected by pine wilt disease in western Japan followed on vegetation maps

Abstract. Succession, changes in the distribution pattern of forest vegetation, and Pinus forest survival following pine wilt disease were clarified based on phytosociological analysis and vegetation maps. Survival of Pinus forests was restricted to the early successional stages, which were located on ridges and the upper part of slopes. Subsequent to pine wilt disease, the succession progressed from early to late substages of Pinus forest, mixed deciduous and evergreen Quercus, to evergreen Quercus forest. Succession occurs in abandoned pine forests which apparently are in a bad state and are vulnerable to attacks by pine wilt disease.     

Dispersal traits as indicators of vegetation dynamics in long-term old-field succession

Succession is a key ecological process that supports our understanding of community assembly and biotic interactions. Dispersal potential and dispersal strategies, such as wind- or animal-dispersal, have been assumed to be highly relevant for the success of plant species during succession. However, research yielded varying results on changes in dispersal modes between successional stages. Here, we test the hypotheses that (a) vascular plant species that use a number of dispersal modes dominate in early stages of succession while species specialized on one/few dispersal modes increase in abundance towards later stages of succession; (b) species well adapted to wind-dispersal (anemochory) will peak in abundance in early successional stages and (c) species well adapted to adhesive dispersal (epizoochory) will increase with proceeding succession. We test these hypotheses in four sites within agriculturally dominated landscapes in Germany. Agricultural use in these sites was abandoned 20–28 years ago, leaving them to secondary succession. Sites have been monitored for plant biodiversity ever since. We analyze changes in plant species richness and abundance, number of dispersal modes and two ranking indices for wind- and adhesive dispersal by applying generalized linear mixed-effect models. We used both abundance-weighted and unweighted dispersal traits in order to gain a comprehensive picture of successional developments. Hypothesis (a) was supported by unweighted but not abundance-weighted data. Anemochory showed no consistent changes across sites. In contrast, epizoochory (especially when not weighted by abundance) turned out to be an indicator of the transition from early to mid-successional stages. It increased for the first 9–16 years of succession but declined afterwards. Species richness showed an opposing pattern, while species abundance increased asymptotically. We suggest that plant-animal interactions play a key role in mediating these processes: By importing seeds of highly competitive plant species, animals are likely to promote the increasing abundance of a few dominant, highly epizoochorous species. These species outcompete weak competitors and species richness decreases. However, animals should as well promote the subsequent increase of species richness by disturbing the sites and creating small open patches. These patches are colonized by weaker competitors that are not necessarily dispersed by animals. The changes in the presence of epizoochorous species indicate the importance of plant traits and related plant–animal interactions in the succession of plant communities.     

Microbial community composition but not diversity changes along succession in arctic sand dunes

The generality of increasing diversity of fungi and bacteria across arctic sand dune succession was tested. Microbial communities were examined by high‐throughput sequencing of 16S rRNA genes (bacteria) and internal transcribed spacer (ITS) regions (fungi). We studied four microbial compartments (inside leaf, inside root, rhizosphere and bulk soil) and characterized microbes associated with a single plant species (Deschampsia flexuosa) across two sand dune successional stages (early and late). Bacterial richness increased across succession in bulk soil and leaf endosphere. In contrast, soil fungal richness remained constant while root endosphere fungal richness increased across succession. There was, however, no significant difference in Shannon diversity indices between early and late successional stage in any compartment. There was a significant difference in the composition of microbial communities between early and late successional stage in all compartments, although the major microbial OTUs were shared between early and late successional stage. Co‐occurrence network analysis revealed successional stage‐specific microbial groups. There were more co‐occurring modules in early successional stage than in late stage. Altogether, these results emphasize that succession strongly affects distribution of microbial species, but not microbial diversity in arctic sand dune ecosystem and that fungi and bacteria may not follow the same successional trajectories.     

Impacts of succession and grass encroachment on heathland Orthoptera

For the conservation of biodiversity, heathlands present important ecosystems throughout Europe. The formerly widespread habitats are nowadays restricted to small and isolated remnants. Without land use heathland vegetation undergoes succession and, in addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses. In the present study we analysed the effects of succession and grass encroachment on Orthoptera in a coastal heathland on the Baltic island of Hiddensee, Germany. Vegetation, microclimate, soil humidity and Orthoptera were sampled in the five main stages of heathland succession, namely grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, and birch forest. Vegetation and environmental parameters showed strong differences among the successional stages. Orthoptera species richness was highest in transitional stages. The high proportion of grasses offer favourable habitat conditions for graminivorous, chorto- and thamnobiont species. Orthoptera density was highest in grey dunes. Threatened and specialised species were restricted to the young stages grey dunes and dwarf-shrub heath. Hence, in order to maintain a high diversity of Orthoptera in heathlands, maintaining different successional stages is of critical importance and this should be integrated into heathland management practices.     

Plant hydraulics and photosynthesis of 34 woody species from different successional stages of subtropical forests

It is important to understand the ecophysiological characters of plants when exploring mechanisms underlying species substitution in the process of plant succession. In the present study, we selected 34 woody species from different stages of secondary succession in subtropical forests of southern China, and measured their hydraulic conductivity, gas exchange rates, leaf nutrients and drought‐tolerance traits such as xylem resistance to cavitation, turgor loss point and carbon isotope ratio. Principal component analysis revealed that early‐, mid‐ and late‐successional species were significantly separated along axis 1, which was strongly associated with hydraulic‐photosynthetic coordination. In contrast to species distributed in late‐successional forest, early‐successional species had the highest hydraulic conductivity, net photosynthetic rates, photosynthetic nitrogen and phosphorus use efficiencies, but had the lowest photosynthetic water‐use efficiency. However, changes of the measured drought‐tolerance traits of the 34 species along the succession did not demonstrate a clear trend – no significant correlations between these traits and plant successional stages were found. Moreover, the trade‐off between hydraulic efficiency and safety was not identified. Taken together, our results suggested that hydraulic efficiency and photosynthetic function, rather than drought tolerance, play an important role in species distributions along plant succession in subtropical forests.     

长白山阔叶红松林不同演替阶段土壤团聚体粒径组成及有机碳含量变化

阔叶红松林是我国东北重要的原生群落,其土壤团聚体在森林生态系统碳固定中具有重要作用.本研究采用空间代替时间的方法,选取白桦幼龄林、白桦中龄林、白桦成熟林、阔叶红松成熟林和阔叶红松过熟林5个不同演替序列,通过湿筛法研究长白山天然针阔混交林群落恢复演替中土壤团聚体粒径组成及有机碳含量的变化.结果表明: 土壤团聚体粒径组成受演替过程影响较大,不同演替阶段下土壤团聚体各粒级所占比例差异显著.团聚体平均质量直径随演替的进行表现为先升高再降低的单峰形式,且最高点出现在白桦成熟林阶段.土壤中不同粒级的团聚体内有机碳含量随着演替的进行呈先增加后略有下降的趋势,且团聚体内有机碳含量最大值出现在阔叶红松成熟林阶段.在同一演替阶段下,0～5和5～10 cm土层(除演替末期的阔叶红松过熟林外)中的各粒径团聚体内有机碳含量都随着粒径的减小而增加,而10～20 cm土层中的各粒径团聚体内有机碳含量都随着粒径的减小而减小.从演替初期的白桦幼龄林到演替末期的阔叶红松过熟林,每个样地内的同一粒径团聚体内有机碳含量均具有明显的垂直分布特性,均随着土层深度的增加而显著降低.