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.

Tree species richness as an important biotic factor regulates the soil phosphorus density in China's mature natural forests.

Tree species richness has been recognized as an underlying driving factor for regulating soil phosphorus (P) status in many site-specific studies. However, it remains poorly understood whether this is true at broad scales where soil P strongly rely on climate, soil type and vegetation type. Here, based on the data of 946 mature natural forest sites from a nationwide field survey in China, we analyzed the impact of tree species richness on soil P density of China's mature natural forests (deciduous coniferous forest, DCF; evergreen coniferous forest, ECF; deciduous broad-leaved forest, DBF; evergreen broad-leaved forest, EBF; and mixed coniferous and broad-leaved forest, MF). Our results showed that tree species richness had a negative effect on soil P density in China's mature natural forests. The Random Forest regression model showed that the relative importance of tree species richness to soil P density was second only to the climate factors (mean annual temperature, MAT; mean annual precipitation, MAP). In addition, the structural equation model (SEM) results showed that the goodness fit of SEM increased when the tree species richness was included into the model. These results suggested that tree species richness was an important factor in regulating the China's mature natural forests soil P density. Furthermore, the SEM results showed that the decreased soil P density was related to the increase in ANPP and the decrease in litter P concentration induced by tree species richness. This result indicates that tree species richness could facilitate plant P absorption and inhibit plant P return into the soil, and thus reducing the soil P density in China's mature natural forests. In conclusion, we found tree species richness was an important biotic factor in regulating soil P density at broad scales, which should be fully considered in Earth models that represent P cycle.

Warming drives sustained plant phosphorus demand in a humid tropical forest.

Phosphorus (P) is often one of the most limiting nutrients in highly weathered soils of humid tropical forests and may regulate the responses of carbon (C) feedback to climate warming. However, the response of P to warming at the ecosystem level in tropical forests is not well understood because previous studies have not comprehensively assessed changes in multiple P processes associated with warming. Here, we detected changes in the ecosystem P cycle in response to a 7-year continuous warming experiment by translocating model plant-soil ecosystems across a 600-m elevation gradient, equivalent to a temperature change of 2.1°C. We found that warming increased plant P content (55.4%) and decreased foliar N:P. Increased plant P content was supplied by multiple processes, including enhanced plant P resorption (9.7%), soil P mineralization (15.5% decrease in moderately available organic P), and dissolution (6.8% decrease in iron-bound inorganic P), without changing litter P mineralization and leachate P. These findings suggest that warming sustained plant P demand by increasing the biological and geochemical controls of the plant-soil P-cycle, which has important implications for C fixation in P-deficient and highly productive tropical forests in future warmer climates.

Natural forests promote phosphorus retention in soil.

Soil phosphorus (P) availability often limits plant productivity. Classical theories suggest that total P content declines at the temporal scale of pedogenesis, and ecosystems develop toward the efficient use of scarce P during succession. However, the trajectory of ecosystem P within shorter time scales of succession remains unclear. We analyzed changes to P pools at the early (I), middle (II), and late (III) stages of growth of plantation forests (PFs) and the successional stages of natural forests (NFs) at 1969 sites in China. We found significantly lower P contents at later growth stages compared to earlier ones in the PF (p < .05), but higher contents at late successional stages than in earlier stages in the NF (p < .05). Our results indicate that increasing P demand of natural vegetation during succession, may raise, retain, and accumulate P from deeper soil layers. In contrast, ecosystem P in PF was depleted by the more rapidly increasing demand outpacing the development of a P-efficient system. We advocate for more studies to illuminate the mechanisms for determining the divergent changes, which would improve forest management and avoid the vast degradation of PF ecosystems suffering from the ongoing depletion of P.

Effects of urbanization on plant phosphorus availability in broadleaf and needleleaf subtropical forests.

Urbanization, the migration of populations from rural to urban areas, has been causing great stress on natural environments, leading to air pollution and nitrogen (N) deposition, negatively affecting forest health. Although there is evidence that urbanization has changed forest N cycling, little is known about whether urbanization also changes the availability of phosphorus (P), which is important for plant growth and forest productivity. To address this question, we carried out a survey in the Pearl River Delta region, the world's largest urban area in southern China, using two types of representative forests, the evergreen broadleaf forests (BFs) and pine plantations (PPs). The leaf N:P ratios in the two forest types were high (20-50) with a significant increasing pattern along the rural-to-urban gradient. The ratios of leaf P:K and P:Na declined along the rural-to-urban gradient, whereas leaf P content did not change in BF but decreased in PP along the rural-to-urban gradient, suggesting that leaf P became limiting along urbanization. The abundance of actinomycetes and gram-negative bacteria decreased along the rural-to-urban gradient, indicating the negative effects of urbanization on soil microorganisms. Principal component analysis indicated that divergent key factors respond to the urbanization and affect plant P limitation in the two forest types. In BF, broadleaf trees showed a greater response to N deposition from urbanization indicating direct leaf N uptake from N deposition is a key factor for plant P limitation. Alternatively, in PP, our findings suggest soil acidification is an important factor accelerating plant P limitation. Our study revealed that urbanization intensifies plant P limitation in subtropical forests, and the effects vary depending on forest types. Our findings provide empirical information to support the management of forest ecosystems and evaluation of urbanization effects on forest health.

Phosphorus uptake in four tree species under nitrogen addition in subtropical China.

Atmospheric N deposition is a serious problem in subtropical China where N is present in large amounts but P is deficient. Several studies hypothesized that N2 fixers can overcome phosphorus limitation by trading fixed N2 for soil phosphorus. In order to know whether N2 fixers could invest fixed N2 in extracellular phosphatase production and could stimulate arbuscular mycorrhizal fungi (AMF) to acquire soil P in N-rich subtropical China, an open-air greenhouse experiment was carried out. Two N2 fixers (Acacia mangium and Ormosia pinnata) and two non-N2 fixers (Schima superba and Pinus massoniana) were exposed to three levels of N addition: 5.6 kg ha-1 a-1 (ambient N), 15.6 kg ha-1 a-1 (middle N), and 20.6 kg ha-1 a-1 (high N). We found that the capacity of plants to acquire soil P in N-rich subtropical China is species specific. The higher P uptake rates were found for N2 fixers than non-N2 fixers under N addition, which were related to the greater soil APA and higher AMF (p < 0.01) in the soil of N2 fixers. However, with time, high N addition decreased more significant quantities of soil microbial phospholipid fatty acids (PLFAs) in the soil of N2 fixers than that of non-N2 fixers (p < 0.05). We conclude that N2 fixers have higher P uptake capacity than non-N2 fixers under ambient N deposition in subtropical China. However, continuing N deposition in the future might affect P uptake ability of N2 fixers as high N addition would decrease soil microbial PLFAs of N2 fixers.

Plant stoichiometric responses to elevated CO2 vary with nitrogen and phosphorus inputs: Evidence from a global-scale meta-analysis.

Rising levels of atmospheric CO2 have been implicated in changes in the nitrogen (N) and phosphorus (P) content of terrestrial vegetation; however, questions remain over the role of C, N and P interactions in driving plant nutrient stoichiometry, particularly whether N and P additions alter vegetation responses to CO2 enrichment singly. Here we use meta-analysis of 46 published studies to investigate the response of plant N and P to elevated CO2 alone and in combination with nutrient (N and P) additions across temperate vs. tropical biomes. Elevated CO2 reduces plant N concentrations more than plant P concentrations in total biomass pools, resulting in a significant decline in vegetation N/P. However, elevated CO2 treatments in combination with N additions increase plant P concentrations, whereas P additions have no statistical effect on plant N concentrations under CO2 enrichment. These results point to compensatory but asymmetrical interactions between N, P and CO2; that changes in N rapidly alter the availability of P, but not the converse, in response to increased CO2. Our finding implies widespread N limitation with increasing atmospheric CO2 concentrations alone. We also suggest that increased anthropogenic N deposition inputs could enhance plant N and P in a progressively CO2-enriched biosphere.

Mineral elements of subtropical tree seedlings in response to elevated carbon dioxide and nitrogen addition.

Mineral elements in plants have been strongly affected by increased atmospheric carbon dioxide (CO2) concentrations and nitrogen (N) deposition due to human activities. However, such understanding is largely limited to N and phosphorus in grassland. Using open-top chambers, we examined the concentrations of potassium (K), calcium (Ca), magnesium (Mg), aluminum (Al), copper (Cu) and manganese (Mn) in the leaves and roots of the seedlings of five subtropical tree species in response to elevated CO2 (ca. 700 µmol CO2 mol(-1)) and N addition (100 kg N ha(-1) yr(-1)) from 2005 to 2009. These mineral elements in the roots responded more strongly to elevated CO2 and N addition than those in the leaves. Elevated CO2 did not consistently decrease the concentrations of plant mineral elements, with increases in K, Al, Cu and Mn in some tree species. N addition decreased K and had no influence on Cu in the five tree species. Given the shifts in plant mineral elements, Schima superba and Castanopsis hystrix were less responsive to elevated CO2 and N addition alone, respectively. Our results indicate that plant stoichiometry would be altered by increasing CO2 and N deposition, and K would likely become a limiting nutrient under increasing N deposition in subtropics.

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.

Nitrogen to phosphorus ratios of tree species in response to elevated carbon dioxide and nitrogen addition in subtropical forests.

Increased atmospheric carbon dioxide (CO2 ) concentrations and nitrogen (N) deposition induced by human activities have greatly influenced the stoichiometry of N and phosphorus (P). We used model forest ecosystems in open-top chambers to study the effects of elevated CO2 (ca. 700 µmol mol(-1) ) alone and together with N addition (100 kg N ha(-1)  yr(-1) ) on N to P (N : P) ratios in leaves, stems and roots of five tree species, including four non-N2 fixers and one N2 fixer, in subtropical China from 2006 to 2009. Elevated CO2 decreased or had no effects on N : P ratios in plant tissues of tree species. N addition, especially under elevated CO2 , lowered N : P ratios in the N2 fixer, and this effect was significant in the stems and the roots. However, only one species of the non-N2 fixers showed significantly lower N : P ratios under N addition in 2009, and the others were not affected by N addition. The reductions of N : P ratios in response to elevated CO2 and N addition were mainly associated with the increases in P concentrations. Our results imply that elevated CO2 and N addition could facilitate tree species to mitigate P limitation by more strongly influencing P dynamics than N in the subtropical forests.

Effects of elevated carbon dioxide and nitrogen addition on foliar stoichiometry of nitrogen and phosphorus of five tree species in subtropical model forest ecosystems.

The effects of elevated carbon dioxide (CO2) and nitrogen (N) addition on foliar N and phosphorus (P) stoichiometry were investigated in five native tree species (four non-N2 fixers and one N2 fixer) in open-top chambers in southern China from 2005 to 2009. The high foliar N:P ratios induced by high foliar N and low foliar P indicate that plants may be more limited by P than by N. The changes in foliar N:P ratios were largely determined by P dynamics rather than N under both elevated CO2 and N addition. Foliar N:P ratios in the non-N2 fixers showed some negative responses to elevated CO2, while N addition reduced foliar N:P ratios in the N2 fixer. The results suggest that N addition would facilitate the N2 fixer rather than the non-N2 fixers to regulate the stoichiometric balance under elevated CO2.

Simulated effects of acidic solutions on element dynamics in monsoon evergreen broad-leaved forest at Dinghushan, China. Part 1: dynamics of K, Na, Ca, Mg and P.

BACKGROUND: Acid deposition has become a concern in south China in recent years. This phenomenon has increased to a dramatic extent with the large use of cars and coal-fueled power plants. As a consequence, soils are becoming acidified and their element dynamics will change. A decrease in the nutrient availability will lead to slower plant growth and maybe to a change in the forest type with current species being replaced by new ones with less nutrient requirements. Because of these reasons, it is important to understand how the dynamics of elements will change and what mechanism is part of the process. This knowledge is important for modeling the acidification process and either finding ways to counter it or to predict its consequences. The primary purpose of this study was to provide information about how the dynamics of K, Na, Ca, Mg and P are affected by acid deposition in a typical forest in southern China. METHODS: Experimental soils and saplings were collected directly from the monsoon evergreen broad-leaved forest in Dinghushan. All saplings were transplanted individually into ceramic pots in August 2000 and placed in an open area near their origin site. Pot soils were treated weekly from October 2000 to July 2002 with an acidic solution at pH 3.05, pH 3.52, pH 4.00 or pH 4.40, or with tap water as a control. The concentrations of SO4(2-), NO3-, K+, Na+, Ca2+, Mg2+ and available P and the pH were measured in soil and leachate samples taken at different times. The sapling leaves were collected and their element concentrations were measured at the end of the experiment. RESULTS AND DISCUSSION: Concentrations of soil exchangeable Ca and Mg decreased quickly over time, although only Ca showed changes with the acidic solution treatment and soil exchangeable K was stable because of soil weathering. Leaching of K, Mg and Ca was dependent upon the treatment acidity. Soil available P decreased slowly without any correlation with the acidity of the treatment. All the NO3- added by the treatment was taken up by the plants, but the SO4(2-) added accumulated in the soil. Amongst the plant species, Schima superba was little affected by the treatment, the leaf P content was affected in Acmena acuminatissima plants and Cryptocarya concinna was the most susceptible species to soil acidification, with a marked decrease of, the leaf K, Ca and Mg concentrations when the treatment acidity increased. CONCLUSIONS: Simulated acid deposition affected the dynamics of K, Ca and Mg in the monsoon evergreen broad-leaved forest. The dynamics of Ca in the soil and of K, Mg and Ca in the soil leachates were affected by the acidic solution treatment. If such a soil acidification occurs, Cryptocarya concinna will be amongst the first affected species, but Schima superba will be able to sustain a good growth and mineral nutrition. RECOMMENDATIONS AND PERSPECTIVES: Acid deposition will lead to imbalance the nutrient elements in the evergreen broad-leaved forest because of accelerated leaching losses of soil exchangeable Ca and Mg. Measures should be developed to slow down soil acidification or nutrient decrease.

[A preliminary study on the chemical properties of precipitation, throughfall, stemflow and surface run-off in major forest types at Dinghushan under acid deposition].

Studies on the chemical properties of precipitation, throughfall, stemflow and surface run-off in major forest types at Dinghushan under acid deposition showed that the pH value of precipitation was about 4.90, and the frequency of acid rain was over 62%. In broad-leaved forest, the pH value of precipitation was lower than that of throughfall, but higher than that of stemflow and especially the surface run-off, indicating that the soil was naturally acidified. In mixed forest, both throughfall and surface run-off had a higher pH value, but stemflow had a lower pH value than precipitation. The throughfall and stemflow were more acidified than precipitation in coniferous pine forest, but the surface run-off had a higher pH value than precipitation. These results suggested that among the three major forest types at Dinghushan, the canopy of broad-leaved forest had the highest buffering ability, whereas for the soil, the coniferous forest had the highest soil buffering capacity. The concentrations of nutrient elements, such as P, K, Ca, Na and Mg in the throughfall, stemflow and surface run-off were higher than those in bulk precipitation in all forests at Dinghushan, some even 10 times higher, indicating that a large amount of nutrients were leached from the canopy. The concentrations of nutrient elements in stemflow were higher than those in throughfall in all forests, and the concentration of nutrient elements in surface water was higher than those in atmospheric rainfall. Coniferous forest had a higher concentration of nutrients in the throughfall and stemflow and a lower nutrient concentration in the surface run-off than other forest types, which implied that nutrient loss was more serious in broad-leaved and mixed forests than in coniferous forests.