Effects of disturbances on the carbon balance of tropical peat swamp forests

Tropical peatlands have accumulated huge soil carbon over millennia. However, the carbon pool is presently disturbed on a large scale by land development and management, and consequently has become vulnerable. Peat degradation occurs most rapidly and massively in Indonesia, because of fires, drainage, and deforestation of swamp forests coexisting with tropical peat. Peat burning releases carbon dioxide (CO₂) intensively but occasionally, whereas drainage increases CO₂ emission steadily through the acceleration of aerobic peat decomposition. Therefore, tropical peatlands present the threat of switching from a carbon sink to a carbon source to the atmosphere. However, the ecosystem‐scale carbon exchange is still not known in tropical peatlands. A long‐term field experiment in Central Kalimantan, Indonesia showed that tropical peat ecosystems, including a relatively intact peat swamp forest with little drainage (UF), a drained swamp forest (DF), and a drained burnt swamp forest (DB), functioned as net carbon sources. Mean annual net ecosystem CO₂ exchange (NEE) (± a standard deviation) for 4 years from July 2004 to July 2008 was 174 ± 203, 328 ± 204 and 499 ± 72 gC m^?2 yr^?1, respectively, for the UF, DF, and DB sites. The carbon emissions increased according to disturbance degrees. We found that the carbon balance of each ecosystem was chiefly controlled by groundwater level (GWL). The NEE showed a linear relationship with GWL on an annual basis. The relationships suggest that annual CO₂ emissions increase by 79–238 gC m^?2 every 0.1 m of GWL lowering probably because of the enhancement of oxidative peat decomposition. In addition, CO₂ uptake by vegetation photosynthesis was reduced by shading due to dense smoke from peat fires ignited accidentally or for agricultural practices. Our results may indicate that tropical peatland ecosystems are no longer a carbon sink under the pressure of human activities.     

Organic matter dynamics control plant species coexistence in a tropical peat swamp forest

We studied the relationship between the coexistence of tree species and the dynamics of organic matter in forests. A tropical peat swamp forest was selected as a model ecosystem, where abiotic factors, such as geological topography or parent rock types, are homogeneous and only biological processes create habitat heterogeneity. The temporal or spatial variation of the ground elevation of peat soils is mainly caused by changes in the balance between organic matter inputs to soils and decomposition, which is affected by the growth and death of influential trees. To clarify the processes of elevation dynamics, we measured the microtopography around some tree groups, estimated organic matter (in the form of litter and roots) in soils under three kinds of microtopographic conditions, measured decomposition rates and detected dominant species' shifting distribution patterns in different stages of growth in relation to the locations of tree groups creating specific microtopographic conditions. We found that growth or death of buttressed trees has the greatest effects on the rising or sinking of ground surfaces through changes in litter supply and root production. We discuss here the possibility of extending our model to other forest types.     

Interdependence of peat and vegetation in a tropical peat swamp forest

The visual uniformity of tropical peat swamp forest masks the considerable variation in forest structure that has evolved in response to differences and changes in peat characteristics over many millennia. Details are presented of forest structure and tree composition of the principal peat swamp forest types in the upper catchment of Sungai Sebangau, Central Kalimantan, Indonesia, in relation to thickness and hydrology of the peat. Consideration is given to data on peat geochemistry and age of peat that provide evidence of the ombrotrophic nature of this vast peatland and its mode of formation. The future sustainability of this ecosystem is predicted from information available on climate change and human impact in this region.     

Surface peat structure and chemistry in a tropical peat swamp forest

Background and Aims

Tropical peat swamp forests (PSF) are great stores of terrestrial carbon and host unique biodiversity. Despite their importance for carbon accounting, the peat characteristics are sparsely studied, and the effect of microtopography on peat properties has not been reported before.

Methods

We compared PSF peat soil characteristics down to 70 cm under differing microtopographical conditions and hydrology.

Results

Long-term water table level data combined with the data from peat structure and chemistry analyses showed differences in most of the measured properties between hummocks and hollows. Decomposition degree was lowest at hummock and hollow surfaces while bulk density and C content increased towards deeper peat. Ash, P, K, Ca and Mg had highest concentrations on hummock surfaces with declining trend downwards, whereas N had no clear concentration pattern along the elevation gradient.

Conclusions

The microtopographical features may not only differ in regards to the water table-induced oxygen conditions but also due to differences in nutrient dynamics.     

Evapotranspiration of tropical peat swamp forests

In Southeast Asia, peatland is widely distributed and has accumulated a massive amount of soil carbon, coexisting with peat swamp forest (PSF). The peatland, however, has been rapidly degraded by deforestation, fires, and drainage for the last two decades. Such disturbances change hydrological conditions, typically groundwater level (GWL), and accelerate oxidative peat decomposition. Evapotranspiration (ET) is a major determinant of GWL, whereas information on the ET of PSF is limited. Therefore, we measured ET using the eddy covariance technique for 4–6 years between 2002 and 2009, including El Niño and La Niña events, at three sites in Central Kalimantan, Indonesia. The sites were different in disturbance degree: a PSF with little drainage (UF), a heavily drained PSF (DF), and a drained burnt ex‐PSF (DB); GWL was significantly lowered at DF, especially in the dry season. The ET showed a clear seasonal variation with a peak in the mid‐dry season and a large decrease in the late dry season, mainly following seasonal variation in net radiation (R_n). The R_n drastically decreased with dense smoke from peat fires in the late dry season. Annual ET forced to close energy balance for 4 years was 1636 ± 53, 1553 ± 117, and 1374 ± 75 mm yr^?1 (mean ± 1 standard deviation), respectively, at UF, DF, and DB. The undrained PSF (UF) had high and rather stable annual ET, independently of El Niño and La Niña events, in comparison with other tropical rainforests. The minimum monthly‐mean GWL explained 80% of interannual variation in ET for the forest sites (UF and DF); the positive relationship between ET and GWL indicates that drainage by a canal decreased ET at DF through lowering GWL. In addition, ET was decreased by 16% at DB in comparison with UF chiefly because of vegetation loss through fires.     

Post‐fire carbon dynamics in the tropical peat swamp forests of Brunei reveal long‐term elevated CH4 flux

Tropical peatlands hold about 15%–19% of the global peat carbon (C) pool of which 77% is stored in the peat swamp forests (PSFs) of Southeast Asia. Nonetheless, these PSFs have been drained, exploited for timber and land for agriculture, leading to frequent fires in the region. The physico‐chemical characteristics of peat, as well as the hydrology of PSFs are affected after a fire, during which the ecosystem can act as a C source for decades, as C emissions to the atmosphere exceed photosynthesis. In this work, we studied the longer‐term impact of fires on C cycling in tropical PSFs, hence we quantified the magnitude and patterns of C loss (CO₂, CH₄ and dissolved organic carbon) and soil‐water quality characteristics in an intact and a degraded burnt PSF in Brunei Darussalam affected by seven fires over the last 40 years. We used natural tracers such as ¹⁴C to investigate the age and sources of C contributing to ecosystem respiration (R_eco) and CH₄, while we continuously monitored soil temperature and water table (WT) level from June 2017 to January 2019. Our results showed a major difference in the physico‐chemical parameters, which in turn affected C dynamics, especially CH₄. Methane effluxes were higher in fire‐affected areas (7.8 ± 2.2 mg CH₄ m^?2 hr^?1) compared to the intact PSF (4.0 ± 2.0 mg CH₄ m^?2 hr^?1) due to prolonged higher WT and more optimal methanogenesis conditions. On the other hand, we did not find significant differences in R_eco between burnt (432 ± 83 mg CO₂ m^?2 hr^?1) and intact PSF (359 ± 76 mg CO₂ m^?2 hr^?1). Radiocarbon analysis showed overall no significant difference between intact and burnt PSF with a modern signature for both CO₂ and CH₄ fluxes implying a microbial preference for the more labile C fraction in the peat matrix.     

Putative roles of bacteria in the carbon and nitrogen cycles in a tropical peat swamp forest

Tropical peat swamp forests are waterlogged, acidic, anoxic and oligotrophic ecosystems. They are important terrestrial carbon pools that help mitigating global warming through carbon sequestration in peat. This study aimed at investigating putative roles of bacteria in the carbon and nitrogen cycles in North Selangor peat swamp forest, Malaysia. Whole-genome sequencing was performed on four bacterial isolates using Illumina NextSeq 500 to decipher their genetic information while Gen III Microplate (Biolog) was applied to verify carbon source utilization. The isolates were identified as Dyella sp. strain C9, Dyella sp. strain C11, Klebsiella sp. strain C31 and Paraburkholderia sp. strain C35. Both Dyella spp. and Paraburkholderia sp. strain C35 were likely novel species while Klebsiella sp. strain C31 was a different strain of the type species, Klebsiella pneumoniae. Both genomic and bioassay results suggested the involvement of the isolates in the degradation of lignocellulose, carbohydrates, sugar alcohols, organic acids and aromatic compounds. The isolates could potentially perform methanotrophy, which helps to mitigate methane emissions from tropical peatlands. In addition, the isolates also contained genes encoding enzymes for nitrite, nitrate and nitric oxide reduction, as well as dissimilatory nitrate reduction to ammonium, which retains the nitrogen in the ecosystems. The results generated insights into potential functions of bacteria in the energy production and nutrient cycling of tropical peatlands, which are essential for the sustainability of high biomass and biodiversity in these ecosystems.     

Forest structure and carbon dynamics in Amazonian tropical rain forests

Living trees constitute one of the major stocks of carbon in tropical forests. A better understanding of variations in the dynamics and structure of tropical forests is necessary for predicting the potential for these ecosystems to lose or store carbon, and for understanding how they recover from disturbance. Amazonian tropical forests occur over a vast area that encompasses differences in topography, climate, and geologic substrate. We observed large differences in forest structure, biomass, and tree growth rates in permanent plots situated in the eastern (near Santarém, Pará), central (near Manaus, Amazonas) and southwestern (near Rio Branco, Acre) Amazon, which differed in dry season length, as well as other factors. Forests at the two sites experiencing longer dry seasons, near Rio Branco and Santarém, had lower stem frequencies (460 and 466 ha^–1 respectively), less biodiversity (Shannon–Wiener diversity index), and smaller aboveground C stocks (140.6 and 122.1 Mg C ha^–1) than the Manaus site (626 trees ha^–1, 180.1 Mg C ha^–1), which had less seasonal variation in rainfall. The forests experiencing longer dry seasons also stored a greater proportion of the total biomass in trees with >50 cm diameter (41–45 vs 30% in Manaus). Rates of annual addition of C to living trees calculated from monthly dendrometer band measurements were 1.9 (Manaus), 2.8 (Santarém), and 2.6 (Rio Branco) Mg C ha^–1 year^–1. At all sites, trees in the 10–30 cm diameter class accounted for the highest proportion of annual growth (38, 55 and 56% in Manaus, Rio Branco and Santarém, respectively). Growth showed marked seasonality, with largest stem diameter increment in the wet season and smallest in the dry season, though this may be confounded by seasonal variation in wood water content. Year-to-year variations in C allocated to stem growth ranged from nearly zero in Rio Branco, to 0.8 Mg C ha^–1 year^–1 in Manaus (40% of annual mean) and 0.9 Mg C ha^–1 year^–1 (33% of annual mean) in Santarém, though this variability showed no significant relation with precipitation among years. Initial estimates of the C balance of live wood including recruitment and mortality as well as growth suggests that live wood biomass is at near steady-state in Manaus, but accumulating at about 1.5 Mg C ha^–1 at the other two sites. The causes of C imbalance in living wood pools in Santarém and Rio Branco sites are unknown, but may be related to previous disturbance at these sites. Based on size distribution and growth rate differences in the three sites, we predict that trees in the Manaus forest have greater mean age (~240 years) than those of the other two forests (~140 years).     

Biomass and carbon dynamics of a tropical mountain rain forest in China

Biometric inventories for 25 years, from 1983 to 2005, indicated that the Jianfengling tropical mountain rain forest in Hainan, China, was either a source or a modest sink of carbon. Overall, this forest was a small carbon sink with an accumulation rate of (0.56±0.22) Mg C ha⁻¹yr⁻¹, integrated from the long-term measurement data of two plots (P9201 and P8302). These findings were similar to those for African and American rain forests ((0.62±0.23) Mg C ha⁻¹yr⁻¹). The carbon density varied between (201.43±29.38) Mg C ha⁻¹ and (229.16±39.2) Mg C ha⁻¹, and averaged (214.17±32.42) Mg C ha⁻¹ for plot P9201. Plot P8302, however, varied between (223.95±45.92) Mg C ha⁻¹ and (254.85±48.86) Mg C ha⁻¹, and averaged (243.35±47.64) Mg C ha⁻¹. Quadratic relationships were found between the strength of carbon sequestration and heavy rainstorms and dry months. Precipitation and evapotranspiration are two major factors controlling carbon sequestration in the tropical mountain rain forest.     

蚂蚁筑巢对西双版纳热带森林土壤碳矿化动态的影响

蚂蚁作为生态系统工程师,能够通过筑巢定居活动增加有机物的输入、改变理化环境及刺激微生物活动,进而影响土壤有机碳矿化动态.本研究以西双版纳高檐蒲桃热带森林群落为研究对象,比较了蚁巢地与非巢地土壤有机碳矿化速率的动态特征,分析蚂蚁筑巢引起的土壤理化性质改变对土壤碳矿化速率的影响.结果表明: 蚂蚁筑巢显著影响土壤有机碳的矿化,相较于非巢地,蚁巢地平均土壤有机碳矿化速率提高19.2%;巢地与非巢地土壤有机碳矿化速率均表现为6月>9月>3月>12月;蚁巢地土壤有机碳矿化速率最大值出现在10～15 cm土层,而非巢地土壤有机碳矿化速率0～5 cm土层最高;蚂蚁筑巢对土壤理化性质产生了显著影响,相较于非蚁巢地,蚁巢地土壤温度、水分、有机碳、微生物生物量碳、全氮、水解氮、硝态氮和铵态氮平均增加幅度分别为7.6%、5.4%、9.9%、14.8%、13.4%、9.9%、24.1%、6.6%和19.4%,而土壤容重和pH平均降幅分别为1.4%和2.5%.相关性分析及主成分分析表明,土壤有机碳和土壤微生物量碳是影响土壤有机碳矿化速率的主控因子,土壤全氮、水解氮、铵态氮、硝态氮、温度和土壤含水率对土壤有机碳矿化的贡献次之.蚂蚁筑巢主要显著改变有机碳矿化的底物组分(土壤有机碳和土壤微生物生物量碳),进而调控西双版纳热带森林土壤有机碳矿化速率的时空动态.     

Land use increases the recalcitrance of tropical peat

Tropical peat carbon compound composition (CCC) is a highly understudied subject. Advanced understanding of peat CCC and carbon dynamics in differing conditions is desperately needed due to large-scale utilization of these peatlands. We studied the CCC—i.e. the hemicellulosic carbohydrate and uronic acid composition and concentrations of extractives, cellulose, acid-soluble lignin and acid-insoluble lignin—in association with peat profile depth and physical structure of peat, under representative, common land uses. Samples were gathered from an undrained forest and three sites altered 20–30 years prior to the study, which in aggregate form a continuum of increasing land-use intensity (drainage-affected forest; drained and deforested degraded open site; drained and deforested site under cultivation) in Central Kalimantan, Indonesia. Peat samples were taken from depths between 10 and 115 cm that covered mostly oxic, frequently waterlogged and permanently waterlogged, anoxic conditions. Our results demonstrated greater modification of peat properties when both vegetation and hydrological conditions were altered. The differences between sites were mainly present in the topmost peat and decreased with depth. Peat located at the surface contained more labile compounds (hemicelluloses, extractives, uronic acids, cellulose) on forest sites than at the most intensively altered open sites, where peat was enriched with recalcitrant acid insoluble lignin. The effect of drainage was evident in the drained forest site, where at the approximate median water table depth peat more closely resembled open sites in terms of the peat properties. The increased recalcitrance of peat in reclaimed areas has been a result of enhanced decomposition, reduced litter input rates and, at open sites also by repeated fires.     

Through enhanced tree dynamics carbon dioxide enrichment may cause tropical forests to lose carbon

The fixation and storage of C by tropical forests, which contain close to half of the globe's biomass C, may be affected by elevated atmospheric CO2 concentration. Classical theoretical approaches assume a uniform stimulation of photosynthesis and growth across taxa. Direct assessments of the C balance either by flux studies or by repeated forest inventories also suggest a current net uptake, although magnitudes sometimes exceed those missing required to balance the global C cycle. Reasons for such discrepancies may lie in the nature of forest dynamics and in differential responses of taxa or plant functional types. In this contribution I argue that CO2 enrichment may cause forests to become more dynamic and that faster tree turnover may in fact convert a stimulatory effect of elevated CO2 on photosynthesis and growth into a long-term net biomass C loss by favouring shorter-lived trees of lower wood density. At the least, this is a scenario that deserves inclusion into long-term projections of the C relations of tropical forests. Species and plant functional type specific responses ('biodiversity effects') and forest dynamics need to be accounted for in projections of future C storage and cycling in tropical forests.     

Morphological changes of rhizobia in peat cultures

Morphological changes that take place in peat cultures of several species of rhizobia were examined. These changes seemed to be associated with enhanced survival of cells in peat and after inoculation onto plastic beads, which were used as a model system for seeds. Cell wall changes, in which the periplasmic space appeared to be occluded with electron-dense material, were observed in Rhizobium sp. strain SU343 and Bradyrhizobium lupini WU425 cells after 7 and 14 days in peat, respectively. Nutrient limitation and low O(2) concentration in peat are suggested to be factors involved in the induction of the morphological changes. Polyhydroxybutyrate reserves, which were present in broth-cultured cells of both species of rhizobia, were mobilized after transfer into peat but did not appear to influence survival after inoculation onto beads. Enhanced expression of an iron-manganese superoxide dismutase was also observed after the cells were transferred into peat. We conclude that cell wall thickening in rhizobia after transfer from broth cultures into peat is an adaptive response for long-term survival under nutrient-limited conditions in peat. Cells with thickened walls may also be more resistant to other types of stress, such as that encountered on a seed surface.     

Forest-to-pasture conversion influences on soil organic carbon dynamics in a tropical deciduous forest

On a global basis, nearly 42% of tropical land area is classified as tropical deciduous forest (TDF) (Murphy and Lugo 1986). Currently, this ecosystem has very high deforestation rates; and its conversion to cattle pasture may result in losses of soil organic matter, decreases in soil fertility, and increases in CO₂ flux to the atmosphere. The soil organic matter turnover rate in a TDF after pasture conversion was estimated in Mexico by determining natural abundances of¹³C. Changes in these values would be induced by vegetation changes from the C₃ (forest) to the C₄ (pasture) photosynthetic pathway. The rate of loss of remnant forest-soil organic matter (fSOM) was 2.9 t ha^–1 year^–1 in 7-year-old pasture and decreased to 0.66 t ha^–1 year^–1 by year 11. For up to 3 years, net fSOM level increased in pastures; this increment can be attributed to decomposition of remnant forest roots. The sand-associated SOM fraction was the most and the silt-associated fraction the least depleted. TDF conversion to pasture results in extremely high rates of loss of remnant fSOM that are higher than any reported for any tropical forest.     

Leaf dynamics,self-shading and carbon gain in seedlings of a tropical pioneer tree

We examined leaf dynamics and leaf age gradients of photosynthetic capacity and nitrogen concentration in seedlings of the tropical pioneer tree, Heliocarpus appendiculatus, grown in a factorial design under controlled conditions with two levels each of nutrients, ambient light (light levels incident above the canopy), and self-shading (the gradient of light levels from upper to lower leaves on the shoot). Correlations among these parameters were examined in order to determine the influence of self-shading, and the regulation of standing leaf numbers, on leaf longevity and its association with leaf photosynthetic capacity. Leaf longevity and the number of leaves on the main shoot were both reduced in high light, while in the low light environment, they were reduced in the steeper self-shading gradient. In high nutrients, leaf longevity was reduced whereas leaf number increased. Leaf initiation rates were higher in the high nutrient treatment but were not influenced by either light treatment. Maximum-light saturated photosynthetic rate, on an area basis, was greater in the high light and nutrient treatments, while the decline in photosynthetic capacity in realtion to leaf position on the shoot was more rapid in high light and in low nutrients. Leaf longevity was negatively correlated among treatments with initial photosynthetic capacity. The leaf position at which photosynthetic capacity was predicted to reach zero was positively correlated with the number of leaves on the shoot, supporting the hypothesis that leaf numbers are regulated by patterns of self-shading. The negative association of longevity and initial photosynthetic capacity apparently arises from different associations among gradients of photosynthetic capacity, leaf numbers and leaf initiation rates in relation to light and nutrient availability. The simultaneous consideration of age and position of leaves illuminates the role of self-shading as an important factor influencing leaf senescence and canopy structure and dynamics.     

Effects of nitrogen additions on above- and belowground carbon dynamics in two tropical forests

Anthropogenic nitrogen (N) deposition is increasing rapidly in tropical regions, adding N to ecosystems that often have high background N availability. Tropical forests play an important role in the global carbon (C) cycle, yet the effects of N deposition on C cycling in these ecosystems are poorly understood. We used a field N-fertilization experiment in lower and upper elevation tropical rain forests in Puerto Rico to explore the responses of above- and belowground C pools to N addition. As expected, tree stem growth and litterfall productivity did not respond to N fertilization in either of these N-rich forests, indicating a lack of N limitation to net primary productivity (NPP). In contrast, soil C concentrations increased significantly with N fertilization in both forests, leading to larger C stocks in fertilized plots. However, different soil C pools responded to N fertilization differently. Labile (low density) soil C fractions and live fine roots declined with fertilization, while mineral-associated soil C increased in both forests. Decreased soil CO₂ fluxes in fertilized plots were correlated with smaller labile soil C pools in the lower elevation forest (R² = 0.65, p < 0.05), and with lower live fine root biomass in the upper elevation forest (R² = 0.90, p < 0.05). Our results indicate that soil C storage is sensitive to N deposition in tropical forests, even where plant productivity is not N-limited. The mineral-associated soil C pool has the potential to respond relatively quickly to N additions, and can drive increases in bulk soil C stocks in tropical forests.     

Predator‐induced changes in dissolved organic carbon dynamics

The fate of dissolved organic carbon (DOC) is partly determined by its availability to microbial degradation. Organisms at upper trophic levels could influence the bioavailability of DOC via cascading effects on primary producers and bacteria. Here we experimentally tested whether the presence of fish in aquatic food webs can indirectly affect the composition of the DOC pool. We found that fish had strong positive effects on phytoplankton biomass that affected the dynamics of DOC composition. Specifically, fish increased protein‐like, algae‐derived DOC mid‐experiment, concurrent with the strongest fish‐induced increase in phytoplankton biomass. Fish also increased bacterial abundance, altered the community composition and diversity of bacteria, and temporarily increased DOC compounds with fluorescence properties indicative of microbially‐reprocessed organic matter. Overall, our experiment revealed that fish can positively influence the substrate (algae‐produced DOC) and the key players (bacteria) of the microbial carbon pump. Consequently, fish could contribute to carbon sequestration by stimulating both the production of bioavailable DOC and the microbial degradation of bioavailable to persistent DOC. We propose this as a novel mechanism whereby the loss of predators from global ecosystems could alter carbon cycling.     

Nitrogen mineralization dynamics in grass monocultures

Although Wedin and Tilman (1990) observed large differences in in situ N mineralization among monocultures of five grass species, the mechanisms responsible were unclear. In this study, we found that the species did not change total soil C or N, and soil C: N ratio (range 12.9–14.1) was only slightly, but significantly, changed after four years. Nor did the species significantly affect the total amount of N mineralized (per g soil N) in year-long aerobic laboratory incubations. However, short-term N mineralization rates in the incubations (day 1–day 17) differed significantly among species and were significantly correlated with annual in situ mineralization. When pool sizes and turnover rates of potentially mineralizable N (N_o) were estimated, the best model treated N_o as two pools: a labile pool, which differed among species in size (N_l, range 2–3% of total N) and rate constant (h, range 0.04–0.26 wk^–1), and a larger recalcitrant pool with a constant mineralization rate across species. The rate constant of the labile pool (h) was highly correlated with annual in situ N mineralization (+0.96). Therefore, plant species need only change the dynamics of a small fraction of soil organic matter, in this case estimated to be less than 3%, to have large effects on overall system N dynamics.