Old-growth Subtropical Forest Research Articles

Soil microbes deal with the nitrogen deposition enhanced phosphorus limitation by shifting community structure in an old-growth subtropical forest

Elevated atmospheric nitrogen (N) deposition potentially enhances the degree of phosphorus (P) limitation in tropical and subtropical forests. However, it remains elusive that how soil microorganisms deal with the N deposition-enhanced P limitation. We collected soils experienced 9 years of manipulative N input at various rates (0, 40, and 80 kg N ha−1 y−1) in an old-growth subtropical natural forest. We measured soil total and available carbon (C), N and P, microbial biomass C, N and P, enzyme activities involved in C, N and P acquisition, microbial community structure, as well as net N and P mineralization. Additionally, we calculated element use efficiency and evaluated microbial homeostasis index. Our findings revealed that N input increased microbial biomass C:P (MBC:P) and N:P (MBN:P) ratios. The homeostasis indexes of MBC:P and MBN:P were 0.68 and 0.75, respectively, indicating stoichiometric flexibility. Interestingly, MBC:P and MBN:P correlated significantly with the fungi:bacteria ratio (F:B), not with N and P use efficiencies, net N and P mineralization, and enzyme C:P (EEAC:P) and N:P (EEAN:P) ratios. Furthermore, EEAC:P and EEAN:P correlated positively with F:B but did not negatively correlate with the C:P and N:P ratios of available resources and microbial biomass. The effects of N deposition on MBC:P, MBN:P and EEAN:P became insignificant when including F:B as a covariate. These findings suggest that microbes flexibly adapted to the N deposition enhanced P limitation by changing microbial community structure, which not only alter microbial biomass C:N:P stoichiometry, but also the enzyme production strategy. In summary, our research advances our understanding of how soil microorganisms deal with the N deposition-enhanced soil P limitation in subtropical forests.

Climate change-related biodiversity fluctuations and composition changes in an old-growth subtropical forest: A 26-yr study

The detection and attribution of biodiversity change is of great scientific interest and central to policy effects aimed at meeting biodiversity targets. Yet, how such a diverse climate scenarios influence forest biodiversity and composition dynamics remains unclear, particularly in high diversity systems of subtropical forests. Here we used data collected from the permanent sample plot spanning 26 years in an old-growth subtropical forest. Combining various climatic events (extreme drought, subsequent drought, warming, and windstorm), we analyzed long-term dynamics in multiple metrics: richness, turnover, density, abundance, reordering and stability. We did not observe consistent and directional trends in species richness under various climatic scenarios. Still, drought and windstorm events either reduced species gains or increased species loss, ultimately increased species turnover. Tree density increased significantly over time as a result of rapid increase in smaller individuals due to mortality in larger trees. Climate events caused rapid changes in dominant populations due to a handful of species undergoing strong increases or declines in abundance over time simultaneously. Species abundance composition underwent significant changes, particularly in the presence of drought and windstorm events. High variance ratio and species synchrony weaken community stability under various climate stress. Our study demonstrates that all processes underlying forest community composition changes often occur simultaneously and are equally affected by climate events, necessitating a holistic approach to quantifying community changes. By recognizing the interconnected nature of these processes, future research should accelerate comprehensive understanding and predicting of how forest vegetation responds to global climate change.

Climate change reshapes plant trait spectrum to explain biomass dynamics in an old-growth subtropical forest

Climate change leads to novel species interactions and continues to reshuffle ecological communities, which significantly declines carbon accumulation rates in mature forests. Still, little is known about the potential influence of multiple global change factors on long-term biomass dynamics and functional trait combinations. We used temporal demographic records spanning 26 years and extensive databases of functional traits to assess how old-growth subtropical forest biomass dynamics respond to various climatic change scenarios (extreme drought, subsequent drought, warming, elevated CO2 concentrations, and windstorm). We found that the initial severe drought, subsequent drought and windstorm events increased biomass loss due to tree mortality, which exceeded the biomass gain produced by survivors and recruits, ultimately resulting in more negative net biomass balances. These drought and windstorm events caused massive biomass loss due to tree mortality that tended towards acquisition species with high hydraulic efficiency, whereas biomass growth from survivors and recruits tended to consist of acquisition species with high hydraulic safety. Compensatory growth in this natural forest provided good explanation for the increase in biomass growth after drought and windstorm events. Notably, these dominant-species transitions reduced carbon storage and residence time, forming a positive carbon-climate feedback loop. Our findings suggest that climate changes could alter functional strategies and cause shifts in new dominant species, which could greatly reduce ecological functions and carbon gains of old-growth subtropical forests.

Effects of Tree Diversity, Functional Composition, and Large Trees on the Aboveground Biomass of an Old-Growth Subtropical Forest in Southern China

Forest aboveground biomass (AGB) plays an important role in regulating the global carbon cycle and is thus an essential component of ecosystem functioning. In the relationships between biodiversity and ecosystem functioning (BEF), studies have shown that many biotic factors (e.g., species, functional traits, and large trees) and abiotic factors have significant impacts on AGB. However, the relative strength of these affecting factors remains unclear. In this study, we analyzed woody plants (diameter at breast height [DBH] ≥ 1 cm) within a 1.6 ha plot in an old-growth subtropical natural forest in southern China. We used structural equation models to test the effects of tree diversity (species, phylogenetic, functional, and size inequality), functional composition, large trees, and environmental factors (topography, soil nutrients, and understory light) on AGB. Our results indicated that size inequality, the community-weighted mean of maximum DBH (CWM_MDBH), and large trees had significant, positive effects on AGB (p < 0.001), while lower soil phosphorus content was found to promote an increase in AGB. Furthermore, large trees, which were mostly composed of dominant tree species, were the main driver of AGB, and the effect of functional composition (e.g., CWM_MDBH) on AGB was substantially reduced by large trees. We argue that the selection effect plays a key role in regulating BEF relationships in subtropical natural forests and conclude that retaining large-diameter trees and dominant species, along with sustaining a complex stand structure, are key measures for improving productivity.

Environmental variation, functional diversity and identity predicting community biomass in an old-growth subtropical broad-leaved forest

The relationship between biodiversity and ecosystem functioning (BEF) has emerged as a central issue in ecosystem ecology. Two broad classes of hypotheses, the mass ratio and niche complementarity effects, are proposed to explain the positive relation between biodiversity and ecosystem functioning in multiple experimental and natural systems. Environmental variation is noted as a key regulator of forest community biomass in natural systems and the under-story tree layer is often overlooked, as the canopy tree species are more likely to contribute to total community biomass. We determined community biomass of over-story, mid-story and under-story tree layers and employed environmental factors, functional diversity metrics of leaf traits to explain the variation in community biomass in an old-growth forest. We found that topographic and edaphic factors were of vital importance in regulating the standing biomass of each tree layer. Although functional diversity displayed positive effects on standing biomass across all tree strata, the standing biomass was more affected by functional identity than diversity in over-story and mid-story tree layers. Across all strata, the most significant correlation between functional diversity and standing biomass in under-story tree layer may suggest strengthened interspecific complementary interactions. Our study confirms that environmental variation is a strong driver of forest community biomass, and the relative importance of the mass ratio and the niche complementarity hypothesis vary across tree strata in explaining diversity effects on standing biomass.

Functional Composition Changes of a Subtropical Monsoon Evergreen Broad-Leaved Forest Under Environmental Change

Long-term studies have revealed that forest species composition was shifting under environment change and disturbance induced by loss of large trees. Yet, few studies explicitly analyzed their impacts on composition concurrently. To learn more about impacts of environment change and disturbance on driving forest community, we investigated shifts in functional composition over past 24 years in an old-growth subtropical forest in southern China. We analyzed nine traits that are mainly related to leaf nutrients, photosynthetic capacity, hydraulic conductivity, and drought tolerance of plants and examined hypotheses: (1) The functional composition change over time was directional instead of random fluctuation, (2) drought-tolerant species increased their abundance under soil dryness, (3) both environmental change and disturbance related to changes of functional composition significantly, and (4) initial trait values of quadrats strongly influenced their subsequent change rates in quadrat level (10 × 10 m). We found that species composition had shifted to favor species with high leaf nutrient content, high photosynthesis rate, high hydraulic conductivity, low water-use efficiency, and high drought tolerance traits, which was due to soil dryness and disturbance. These two factors explained 47–58% of quadrats’ trait value changes together. Considering rapidly increasing stem density, this pattern may indicate ecological processes of which disturbance provided numerous recruits of resource-acquisition strategy species and soil dryness conducted a selecting effect on shaping composition in the forest. Additionally, quadrats with initial trait values at the far end of change direction shifted faster in three traits, which also indicated that functional composition changes in quadrats were directional and homogenized. Our results implied that environment change and accompanied disturbance events possibly drove species composition change along a different trajectory in the subtropical forest that experienced high climatic variability.

Species Differences in Nitrogen Acquisition in Humid Subtropical Forest Inferred From 15N Natural Abundance and Its Response to Tracer Addition

Differences in nitrogen (N) acquisition patterns between plant species are often reflected in the natural 15N isotope ratios (δ15N) of the plant tissues, however, such differences are poorly understood for co-occurring plants in tropical and subtropical forests. To evaluate species variation in N acquisition traits, we measured leaf N concentration (%N) and δ15N in tree and understory plant species under ambient N deposition (control) and after a decade of N addition at 50 kg N ha−1 yr−1 (N-plots) in an old-growth subtropical forest in southern China. We also measured changes in leaf δ15N after one-year of 15N addition in both the control and N-plots. The results show consistent significant species variation in leaf %N in both control and N-plots, but decadal N addition did not significantly affect leaf %N. Leaf δ15N values were also significantly different among the plant species both in tree and understory layers, and both in control and N-plots, suggesting differences in N acquisition strategies such as variation in N sources and dominant forms of N uptake and dependence on mycorrhizal associations among the co-occurring plant species. Significant differences between the plant species (in both control and N-plots) in changes in leaf δ15N after 15N addition were observed only in the understory plants, indicating difference in access (or use) of deposited N among the plants. Decadal N addition had species-dependent effects on leaf δ15N, suggesting the N acquisition patterns of these plant species are differently affected by N deposition. These results suggest that co-occurring plants in N-rich and subtropical forests vary in their N acquisition traits; these differences need to be accounted for when evaluating the impact of N deposition on N cycling in these ecosystems.

Canopy openness and topographic habitat drive tree seedling recruitment after snow damage in an old-growth subtropical forest

We investigated the relative and combined effect of topography and light environment on the recruitment of seedlings in a subtropical forest after snow damage to the canopy. The tree seedling community in an old-growth subtropical forest was monitored using 500 2 m × 2 m seedling plots at six-month intervals for 2 years. With a focus on recruitment following canopy damage, we related abiotic and biotic environmental variables to seedling dynamics, and we tested if significant topographic and light habitat associations were present for seedlings recruiting via a torus translation test. Then, we used variance partitioning to examine the relative effects of spatial, topographic and light variables on the temporal assemblages of seedlings. A total of 3047 seedlings from 58 species recruited in the first 2 years following snow damage. At the community level, increases in seedling abundance and richness were positively correlated with canopy openness and negatively correlated with elevation. At the species level, both pioneer and late-successional tree species had more recruits in high light environment than in low light environment. 84.3% of the recruiting seedlings were significantly associated to either light environment (35.7%), topography (26.5%) or both (22.1%). Despite this, at the plot level, spatial variables (PCNM) explained the majority of the variability in seedling composition over time. Our results suggest that snow damage to the canopy increases species richness and abundance via light-facilitated seedling recruitment, and that the composition of recruiting seedlings was largely spatially dependent. Topographic habitat filtering acts as a persistent force in determining the recruitment of seedlings and increases in strength with increased light-facilitated seedling recruitment. Our results highlight that, in this subtropical forest, both light requirement and topographic specialization interact over time to play a key role in promoting coexistence of tree species through selection of individuals at the seedling stage. We also suggest exploring the possibilities of management intervention to speed up the recovery of this forest.

Exogenous Nitrogen Addition Reduced the Temperature Sensitivity of Microbial Respiration without Altering the Microbial Community Composition.

Atmospheric nitrogen (N) deposition is changing in both load quantity and chemical composition. The load effects have been studied extensively, whereas the composition effects remain poorly understood. We conducted a microcosm experiment to study how N chemistry affected the soil microbial community composition characterized by phospholipid fatty acids (PLFAs) and activity indicated by microbial CO2 release. Surface and subsurface soils collected from an old-growth subtropical forest were supplemented with three N-containing materials (ammonium, nitrate, and urea) at the current regional deposition load (50 kg ha-1 yr-1) and incubated at three temperatures (10, 20, and 30°C) to detect the interactive effects of N deposition and temperature. The results showed that the additions of N, regardless of form, did not alter the microbial PLFAs at any of the three temperatures. However, the addition of urea significantly stimulated soil CO2 release in the early incubation stage. Compared with the control, N addition consistently reduced the temperature dependency of microbial respiration, implying that N deposition could potentially weaken the positive feedback of the warming-stimulated soil CO2 release to the atmosphere. The consistent N effects for the surface and subsurface soils suggest that the effects of N on soil microbial communities may be independent of soil chemical contents and stoichiometry.

Paradox Between Species Diversity and Conservation

One way to draw attention to an area regarding conservation is to declare it as an Atlantic Forest Biosphere Reserve (RBMA). In this article, we analyzed attributes related to different forest remnants, including the only RBMA of Santa Catarina state (Brazil), a modified old-growth Subtropical Atlantic Forest remnant. We brought evidences that patterns of distribution of species abundance and species diversity are influenced by highly dominant species. We found a relevant proportion of endemic tree/shrub species and three endangered species. These findings demonstrate the relevant biodiversity of the Subtropical Atlantic Forest of Santa Catarina. The diversity profiles of the RBMA and other areas overlapped. This finding, which has an important conservational implication that the tree species diversity in these areas is similar. Therefore, we concluded that the areas considered in our study deserve attention regarding biological conservation. New priority areas for conservation are necessary, and the esta...

Spatial Heterogeneity of Soil Chemical Properties in a Subtropical Karst Forest, Southwest China

This study evaluates the spatial heterogeneity of the soil chemical properties of surface soils across a 1 ha old-growth subtropical karst forest in southwest China.

Rapid structural and compositional change in an old-growth subtropical forest: using plant traits to identify probable drivers.

Recent studies have shown directional changes in old-growth tropical forests, but changes are complex and diverse, and their drivers unclear. Here, we report rapid net structural and compositional changes in an old-growth subtropical forest and we assess the functional nature of these changes to test hypothetical drivers including recovery from past disturbances, reduction in ungulate browsing, CO2 fertilization, and increases in rainfall and temperature. The study relies on 15 years of demographic monitoring within 8 ha of subtropical montane forest in Argentina. Between 1992 and 2007, stem density markedly increased by 50% (12 stems ha−1 y−1) and basal area by 6% (0.13 m2 ha−1 y−1). Increased stem density resulted from enhanced recruitment of understory treelets (Piper tucumanum, Eugenia uniflora, Allophylus edulis) into small size classes. Among 27 common tree species, net population growth was negatively correlated with maximum tree size and longevity, and positively correlated with leaf size and leaf nutrient content, especially so when initial population size was controlled for. Changes were inconsistent with predictions derived from past disturbances (no increase in shade-tolerant or long-lived late-succesional species), rainfall or temperature increase (no increase in evergreen or deciduous species, respectively). However, the increase in nutrient-rich soft-leaved species was consistent with exclusion of large herbivores two decades before monitoring started; and CO2 fertilization could help explain the disproportionate increase in small stems. Reductions in populations of large vertebrates have been observed in many otherwise undisturbed tropical forests, and our results suggest they can have important structural and functional repercussions in these forests.

Soil respiration in an old‐growth subtropical forest: Patterns, components, and controls

AbstractThe patterns, components, and controls of soil respiration in an old‐growth subtropical forest were investigated using an automatic chamber system. We measured soil respiration in three treatments (control, trenching, litter removal) over 15 months. The annual total soil respiration (1248 gC m–2 yr–1) showed considerable spatial variation (coefficient of variation = 27.8%) within the forest. Thirty samples were required to obtain results within 10% of the mean value at a 95% confidential level. A distinctive cosine‐like diel pattern of soil respiration was observed; the time lag between gross primary production and soil respiration at this scale was calculated to be 4–5 h. Seasonality of soil respiration was strong (~1 µmol m–2 s–1 near the end of winter; ~6 µmol m–2 s–1 in midsummer). No time lag was discerned between gross primary production and soil respiration at the seasonal scale. Soil temperature at 5 cm below surface can explain most (>91%) of the observed annual variation in soil respiration. The apparent respiration temperature sensitivity index (Q10) was 3.05. The lowest Q10 value was observed in winter, when soil moisture was low. Soil respiration was overestimated by a Q10 function during both dry and wet periods. The relative contributions of soil organic matter (RSOM), litterfall decomposition (RL), and root respiration (RR) to total soil respiration are 65.25%, 18.73%, and 16.01%, respectively; the temperature sensitivity of these components differ: RL (Q10 = 7.22) > RSOM (2.73) > RR (1.65). This relationship between Q10 values for litter respiration, soil organic matter decomposition, and root respiration still holds after minimizing the confounding effect of moisture. A relatively constant substrate supply and/or thermal acclimation could account for the observed low‐temperature sensitivity in root respiration. Given the high carbon stocks and fluxes, the old‐growth subtropical forests of China seem important in the global carbon budget and climate change.

Substantial amounts of carbon are sequestered during dry periods in an old-growth subtropical forest in South China

A number of continuous eddy covariance measurements and long-term biomass inventories had proved that old-growth forests are carbon sinks worldwide. The present study estimated the net ecosystem productivity (NEP) for an old-growth subtropical forest at the Dinghushan Biosphere Reserve in South China to investigate the temporal pattern of carbon sequestration, both seasonally and annually. The measured NEP over 7 years (from 2003 to 2009) showed that this forest was a net carbon sink, ranging from 230 (in 2008) to 489…−2 year−1 (in 2004). The greatest value of NEP was found in the driest year and the lowest value in the wettest year during the study period. Within a year, NEP during the dry season was about 81.4 % higher than for the wet season. Accordingly, the dry season at seasonal scale and dry years at interannual scale are key periods for carbon sequestration in this forest. The strong seasonality of ecosystem or soil respiration (ER or SR) compared with gross primary productivity (GPP) resulted in substantial amounts of carbon being sequestered during dry seasons. A decrease of GPP and an increase of ER or SR demonstrated the lower carbon uptake in rainy years. From this study, we conclude that GPP and living biomass carbon increment are not overriding parameters controlling NEP. The variations in ER or SR driven by the rainfall scheme were the dominant factor determining the magnitude of NEP in this forest in South China.

Aggregated spatial distributions of species in a subtropical karst forest, southwestern China

Spatial distribution patterns of species reflect not only the ecological processes but also the habitat features that are related to species distribution. In karst topography, species distribution patterns provide more specific information about their environments. The objectives of this study are as follows: (i) to analyse and explain the spatial distribution patterns of conspecific trees in an old-growth subtropical karst forest; (ii) to investigate pattern changes at different spatial scales; (iii) to test the spatial pattern similarity (or dissimilarity) between trees at different abundances, diameter at breast height classes, canopy layers and different functional groups (shade tolerance and seed dispersal mode); (iv) to examine whether habitat heterogeneity has an important effect on the species spatial distribution. The spatial distributions of woody species with ≥20 individuals in a 1-ha subtropical karst forest plot at Maolan in southwestern China were quantified using the relative neighbourhood density Ω based on the average density of conspecific species in a circular neighbourhood around each species. Aggregated distribution is the dominant pattern in the karst forest, but the ratio of aggregated species in total species number decreases with an increase in spatial scale. Less abundant species are more aggregated than most abundant species. Aggregation is weaker in larger diameter classes, which is consistent with the prediction of self-thinning. Seed dispersal mode influences spatial patterns, with species dispersed by animals being less aggregated than those dispersed by wind and gravity. Other species functional traits (e.g. shade tolerance) also influence the species spatial distributions. Moreover, differences among species habitat associations, e.g. with rocky outcrops, play a significant role in species spatial distributions. These results indicate that habitat heterogeneity, seed dispersal limitation and self-thinning primarily contribute to the species spatial distributions in this subtropical karst forest.

Different patterns of ecosystem carbon accumulation between a young and an old-growth subtropical forest in Southern China

Using long-term (22 years) measurements from a young and an old-growth subtropical forest in southern China, we found that both forests accumulated carbon from 1982 to 2004, with the mean carbon accumulation rate at 227 ± 59 g C m−2 year−1 for young forest and 115 ± 89 g C m−2 year−1 for the old-growth forest. Allocation of the accumulated carbon was quite different between these two forests: the young forest accumulated a significant amount of carbon in plant live biomass, whereas the old-growth forest accumulated a significant amount of carbon in the soil. From 1982 to 2004, net primary productivity (NPP) increased for the young forest, and did not change significantly for the old-growth forest. The increase in NPP of the young forest resulted from recruitment of some dominant tree species characteristic of the subtropical mature forest in the region and an increase in tree density; decline of NPP of the old-growth forest was caused by increased mortality of the dominant trees.