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.     

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.     

Carbon fluxes from a tropical peat swamp forest floor

A tropical ombrotrophic peatland ecosystem is one of the largest terrestrial carbon stores. Flux rates of carbon dioxide (CO₂) and methane (CH₄) were studied at various peat water table depths in a mixed‐type peat swamp forest floor in Central Kalimantan, Indonesia. Temporary gas fluxes on microtopographically differing hummock and hollow peat surfaces were combined with peat water table data to produce annual cumulative flux estimates. Hummocks formed mainly from living and dead tree roots and decaying debris maintained a relatively steady CO₂ emission rate regardless of the water table position in peat. In nearly vegetation‐free hollows, CO₂ emission rates were progressively smaller as the water table rose towards the peat surface. Methane emissions from the peat surface remained small and were detected only in water‐saturated peat. By applying long‐term peat water table data, annual gas emissions from the peat swamp forest floor were estimated to be 3493±316 g CO₂ m^?2 and less than 1.36±0.57 g CH₄ m^?2. On the basis of the carbon emitted, CO₂ is clearly a more important greenhouse gas than CH₄. CO₂ emissions from peat are the highest during the dry season, when the oxic peat layer is at its thickest because of water table lowering.     

Leaf litter decomposition in a tropical peat swamp forest in Peninsular Malaysia

It has long been assumed that the peat underlying tropical peat swamp forests accumulates because the extreme conditions (water logged, nutrient poor, anaerobic and acidic—pH 2.9–3.5) impede microbial activity. Litterbag studies in a tropical Malaysian peat swamp (North Selangor peat swamp forest) showed that although the sclerophyllous, toxic leaves of endemic peat forest plants (Macaranga pruinosa, Campnosperma coriaceum, Pandanus atrocarpus, Stenochlaena palustris) were barely decomposed by bacteria and fungi (decay rates of only 0.0006–0.0016 k day⁻¹), leaves of M. tanarius, a secondary forest species were almost completely decomposed (decay rates of 0.0047–0.005 k day⁻¹) after 1 year. Thus it is intrinsic properties of the leaves (that are adaptations to deter herbivory in the nutrient poor environment) that impede microbial breakdown. The water of the peat swamp was very high in dissolved organic carbon (70–84 mg l⁻¹ DOC). Laboratory studies revealed initial rapid leaching of DOC from leaves (up to 1,720 mg l⁻¹ from 4 g of leaves in 7 days), but the DOC levels then fell rapidly. The leaching of DOC resulted in weight loss but the physical structure of the leaves remained intact. It is suggested that the DOC is used as a substrate for microbial growth hence lowering the concentration of DOC in the water and transferring energy from the leaves to other trophic levels. This would explain how nutrient poor tropical peatswamps support diverse, abundant flora and fauna despite low nutrient levels and lack of rapid litter cycling such as occurs in other types of tropical rainforests.     

Carbon dioxide balance of a tropical peat swamp forest in Kalimantan, Indonesia

Tropical peatlands, which coexist with swamp forests, have accumulated vast amounts of carbon as soil organic matter. Since the 1970s, however, deforestation and drainage have progressed on an enormous scale. In addition, El Niño and Southern Oscillation (ENSO) drought and large‐scale fires, which grow larger under the drought condition, are accelerating peatland devastation. That devastation enhances decomposition of soil organic matter and increases the carbon release to the atmosphere as CO₂. This phenomenon suggests that tropical peatlands have already become a large CO₂ source, but related quantitative information is limited. Therefore, we evaluated the CO₂ balance of a tropical peat swamp forest in Central Kalimantan, Indonesia, using 3 years of CO₂ fluxes measured using the eddy covariance technique from 2002 through 2004. The forest was disturbed by drainage; consequently, groundwater level (GL) was reduced. The net ecosystem CO₂ production (NEP) measurements showed seasonal variation, which was slightly positive or almost zero in the early dry season, and most‐negative late in the dry season or early the rainy season. This seasonality is attributable to the seasonal pattern of climate, tree phenology and fires. Slightly positive NEP resulted from smaller ecosystem respiration (RE) and larger gross primary production (GPP) under conditions of high photosynthetic photon flux density (PPFD) and large leaf area index (LAI). The most‐negative NEP resulted from smaller GPP and larger RE. The smaller GPP was related to high vapor pressure deficit (VPD), small LAI and low PPFD because of smoke from fires. The larger RE was related to low GL. Annual NEP values were estimated respectively as −602, −382 and −313 g C m⁻² yr⁻¹ for 2002, 2003 and 2004. These negative NEP values show that the tropical peat swamp forest, disturbed by drainage, functioned as a CO₂ source. That source intensity was highest in 2002, an ENSO year, mainly because of low PPFD caused by dense smoke emitted from large fires.     

The contribution of leaching to nutrient release from leaf litter of two emergent tree species in a Malaysian tropical peat swamp forest

Ostracode faunas changed in Lake Bonneville as the lake rose in the closed basin and became more dilute, then overflowed, and eventually returned to closed-basin hydrology; alkalinity (carbonate + bicarbonate) increased throughout the lake history. Fossil-ostracode faunas in marl deposited in the lake between about 30 and 13 cal ka allow reconstruction of these geochemical changes (as shown by R. M. Forester), and permit stratigraphic correlations within the large lake basin, including into the Sevier subbasin, which was connected to the main body of the lake but had different inflowing rivers. The first ostracode to appear in Lake Bonneville deposits at low altitudes is Limnocythere staplini; it was replaced by L. ceriotuberosa as the lake got larger and the water became more dilute and alkaline. Candona adunca and several species of Fabaeformiscandona mark the freshest-water phases. Cytherissa lacustris appeared in lake-margin settings after lake level dropped and cold, fresh groundwater began to discharge into the lake. Candona sp. cf. eriensis lived in the Sevier subbasin during the transgressive phase of the lake, but did not appear in the main body until late in the regressive phase. Limnocythere sappaensis, which requires high alkalinity, showed up late in the regressive phase in the main body when lake level was very low.     

Tree species diversity and forest structure in relation to microtopography in a tropical freshwater swamp forest in French Guiana

Diversity of tree association and forest structure were analysed in relation to microtopography and flooding intensity in a tropical freshwater swamp forest in the Sinnamary river basin, French Guiana. A 530-m-long vegetation transect was established through a hummock-hollow terrain. Nine 10 m× 50 m sample plots, perpendicular to the main transect, were located so that each was as microtopographically uniform as possible. Trees with dbh (diameter at 1.3 m) 10 cm were censused in all plots and trees with 2 cm dbh < 10 cm in three plots. Sixty tree species belonging to 39 genera and 30 families were recorded. The study area was divided into low and high sites according to microtopography and flooding intensity. According to the Czekanowski similarity matrix, the tree association in low, most frequently flooded, sites differed from that in the high sites under intermediate or low flooding intensity. The low sites had higher stand density and lower species richness than the high sites. Trees with dbh 10 cm in low sites were small and stand basal area (SBA) was about the same in low (69.6 m² ha^–1) and high (64.3 m² ha^–1) sites. The low areas were dominated by Pterocarpus officinalis (38% of stems with dbh 10 cm and 36% of SBA) and Malouetia tamaquarina (26 and 15%). Diospyros guianensis (13.4% of stems with dbh 10 cm and 6.1% of SBA), a Caraipa sp. (14.0 and 7.9%), Lecythis corrugata (6.6 and 3.5%) and emergent Caryocar microcarpum (0.9 and 13.9%) were abundant in high sites. Nodulated legume trees, P. officinalis, Hydrochorea corymbosa and Inga disticha, comprised 44% of stems in the low sites. The abundant nodulation suggests that symbiotic dinitrogen fixation may be an adaptation to N-depleted waterlogged soils. Other adaptive responses were litter accumulation between the buttresses of P. officinalis, which formed hummocks above surface water, and clonal growth habit of M. tamaquarina, which resulted in formation of monospecific groves in low sites.     

Yeasts from peat in a tropical peat swamp forest in Thailand and their ability to produce ethanol,indole-3-acetic acid and extracellular enzymes

This study aimed to isolate and identify yeasts from peat in To Daeng peat swamp forest in southern of Thailand, and to investigate their ability to produce ethanol from glucose and xylose and to produce indole-3-acetic acid (IAA) and extracellular enzymes. A total of 65 yeast strains were obtained from 15 peat samples using an enrichment technique, and 61 strains were identified to be five species belonging to the phylum Ascomycota, namely Cyberlindnera subsufficiens, Debaryomyces fabryi, Meyerozyma guilliermondii, Saturnispora diversa and Schwanniomyces polymorphus var. africanus, and five species of the phylum Basidiomycota, namely Cryptococcus taiwanensis pro tem, Cutaneotrichosporon mucoides, Papiliotrema flavescens, Papiliotrema laurentii and Rhodotorula mucilaginosa. Four strains were unidentified and require further analysis. They differed from the type strain of P. flavescens by two nucleotide substitutions in the D1/D2 region of the LSU rRNA gene and nine nucleotide substitutions in the ITS region. R. mucilaginosa was the most prevalent yeast species, followed by S. polymorphus var. africanus, Cy. subsufficiens and D. fabryi. None of the yeast strains obtained in this study were able to ferment xylose to ethanol, but all ascomycetous yeast strains produced ethanol from glucose in a range of 9.0–58.0 g/L, with Cy. subsufficiens DMKU-YNB42-1 producing the highest ethanol concentration. A total of 62 strains produced IAA in a range of 9.0 to 66.9 mg/L, with the highest IAA produced by R. mucilaginosa DMKU-Y33-A. Investigation of the production of cellulases, xylanase, pectinase, amylase, protease and lipase revealed that all 65 yeast strains produced at least one extracellular enzyme, a lipase.     

A comparison of sites suitable for the seedling establishment of two co-occurring species, Swintonia glauca and Stemonurus scorpioides, in a tropical peat swamp forest

Spatial and temporal ground-surface dynamics are major factors that affect regeneration and species coexistence in tropical peat swamp forests. We studied the seedling survivorship and morphological features of two tree species that play important roles in maintaining the ground-surface dynamics of a peat swamp forest in Sumatra. Large Swintonia glauca trees form mounds, whereas large Stemonurus scorpioides trees occupy non-mounds. We monitored the demography of naturally dispersed Swintonia and Stemonurus seedlings that germinated in 2000. Survivorship of Swintonia seedlings was high under conditions of late germination, high-light environment, and elevated ground surface, and was negatively affected by distance to the nearest conspecific adult. Survivorship of Stemonurus was high under conditions of early germination and high conspecific seedling density, and was also negatively affected by distance to the nearest conspecific adult. The allometric features of Stemonurus seedlings indicated characteristics of stress tolerance, that is, low growth rate and thick, porous roots. Stemonurus, which has large wingless seeds, regenerated in non-mounds around the parental trees, while winged Swintonia seeds dispersed farther from the parent and established in patchily distributed gaps and mounds. Thus, Swintonia seedlings can survive on non-mound sites within gaps and possibly create mounds, while Stemonurus seedlings tend to maintain non-mounds around the parental trees.     

Plant species coexistence at local scale in temperate swamp forest: test of habitat heterogeneity hypothesis

It has been suggested that a heterogeneous environment enhances species richness and allows for the coexistence of species. However, there is increasing evidence that environmental heterogeneity can have no effect or even a negative effect on plant species richness and plant coexistence at a local scale. We examined whether plant species richness increases with local heterogeneity in the water table depth, microtopography, pH and light availability in a swamp forest community at three local spatial scales (grain: 0.6, 1.2 and 11.4 m). We also used the variance partitioning approach to assess the relative contributions of niche-based and other spatial processes to species occurrence. We found that heterogeneity in microtopography and light availability positively correlated with species richness, in accordance with the habitat heterogeneity hypothesis. However, we recorded different heterogeneity-diversity relationships for particular functional species groups. An increase in the richness of bryophytes and woody plant species was generally related to habitat heterogeneity at all measured spatial scales, whereas a low impact on herbaceous species richness was recorded only at the 11.4 m scale. The distribution of herbaceous plants was primarily explained by other spatial processes, such as dispersal, in contrast to the occurrence of bryophytes, which was better explained by environmental factors. Our results suggest that both niche-based and other spatial processes are important determinants of the plant composition and species turnover at local spatial scales in swamp forests.     

Biogeochemical properties of a tropical swamp forest ecosystem in southern Thailand

The distributions of organic matter in the tropical swamps in southern Thailand are reported. The concentrations of particulate and dissolved organic carbon (POC and DOC) in the Bang Nara River, which drains swamp forests and nearby paddy fields, were 2.9 ± 2.0 and 6.2 ± 1.3 mg C l⁻¹, respectively. Although the variation was large, DOC concentration in the Bang Nara River seemed to be higher than POC in November 1992 (DOC/POC ratio, 2.8 ± 2.2). River waters from the upland areas were characterized by low POC and DOC concentrations as compared with Bang Nara River water. The δ¹³C values of POC and river sediments were useful to distinguish between organic matter originating in upland and swamp areas. It is suggested that the distributions of organic matter and its isotopic composition reflect the difference in drainage characteristics between lowland swamp and upland areas. Isotopic analyses of plant leaves and soils revealed that the swamp forest ecosystems were characterized by low δ¹³C and low δ¹⁵N values, which suggested low efficiency of water use by plants and large contributions of atmospheric deposition of nitrogen, respectively. Although CO₂ recycling in the forest might be an important factor determining the δ¹³C values of understory plants, the main process in carbon metabolism of tropical swamp forests would be CO₂ exchange between the atmosphere and forest canopy. Received: May 1, 2001 / Accepted: September 28, 2001     

Organic matter inputs shift soil enzyme activity and allocation patterns in a wet tropical forest

Soil extracellular enzymes mediate organic matter turnover and nutrient cycling yet remain little studied in one of Earth’s most rapidly changing, productive biomes: tropical forests. Using a long-term leaf litter and throughfall manipulation, we explored relationships between organic matter (OM) inputs, soil chemical properties and enzyme activities in a lowland tropical forest. We assayed six hydrolytic soil enzymes responsible for liberating carbon (C), nitrogen (N) and phosphorus (P), calculated enzyme activities and ratios in control plots versus treatments, and related these to soil biogeochemical variables. While leaf litter addition and removal tended to increase and decrease enzyme activities per gram soil, respectively, shifts in enzyme allocation patterns implied changes in relative nutrient constraints with altered OM inputs. Enzyme activity ratios in control plots suggested strong belowground P constraints; this was exacerbated when litter inputs were curtailed. Conversely, with double litter inputs, increased enzymatic investment in N acquisition indicated elevated N demand. Across all treatments, total soil C correlated more strongly with enzyme activities than soluble C fluxes, and enzyme ratios were sensitive to resource stoichiometry (soil C:N) and N availability (net N mineralization). Despite high annual precipitation in this site (MAP ~5 m), soil moisture positively correlated with five of six enzymes. Our results suggest resource availability regulates tropical soil enzyme activities, soil moisture plays an additional role even in very wet forests, and relative investment in C, N and P degrading enzymes in tropical soils will often be distinct from higher latitude ecosystems yet is sensitive to OM inputs.     

Non-woody life-form contribution to vascular plant species richness in a tropical American forest

We provide total vascular plant species counts for three 1-ha plots in deciduous, semi-deciduous and evergreen forests in central Bolivia. Species richness ranged from 297 species and 22,360 individuals/ha in the dry deciduous forest to 382 species and 31,670 individuals/ha in the evergreen forest. Orchidaceae, Pteridophyta and Leguminosae were among the most species-rich major plant groups in each plot, and Peperomia (Piperaceae), Pleurothallis (Orchidaceae) and Tillandsia (Bromeliaceae), all epiphytes, were the most species-rich genera. This dominance of a few but very diverse and/or widespread taxa contrasted with the low compositional similarity between plots. In a neotropical context, these Central Bolivian forest plots are similar in total species richness to other dry deciduous and humid montane forests, but less rich than most Amazonian forests. Nevertheless, lianas, terrestrial herbs and especially epiphytes proved to be of equal or higher species richness than most other neotropical forest inventories from which data are available. We therefore highlight the importance of non-woody life-forms (especially epiphytes and terrestrial herbs) in Andean foothill forest ecosystems in terms of species richness and numbers of individuals, representing in some cases nearly 50% of the species and more than 75% of the individuals. These figures stress the need for an increased inventory effort on non-woody plant groups in order to accurately direct conservation actions. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Adaptive significance of vegetative sprouting for a tropical canopy tree, Scaphium longiflorum (Sterculiaceae), in a peat swamp forest in Central Kalimantan

Scaphium longiflorum Ridley (Sterculiaceae), a common canopy tree in peat swamp forests in the Far East, produces vegetative sprouting in its juvenile stage. We investigated morphological features and allometric properties of the species in a peat swamp forest in Central Kalimantan, Indonesia, to determine under what conditions ramets are produced and discuss the adaptive significance of the vegetative sprouting in a peat swamp environment. Larger juveniles were more toppled, and the toppled ones sprouted vegetatively. Therefore, the vegetative sprouting acts as a countermeasure for a shoots mechanical failure and fall on an unstable peat soil. We propose three hypotheses to explain the reason why larger juveniles are more often toppled in a peat swamp environment.     

Organic matter chemistry and dynamics in clear-cut and unmanaged hardwood forest ecosystems

Forest harvesting alters the organic matter cycle by changing litter inputs and the decomposition regime. We hypothesized that these changes would result in differences in organic matter chemistry between clear-cut and uncut watershed ecosystems. We studied the chemistry of soil organic matter (SOM), and dissolved organic carbon (DOC) in soil solutions and stream samples in clear-cut and uncut sites at the Hubbard Brook Experimental Forest in New Hampshire using DOC fractionation techniques and solid-state ¹³C nuclear magnetic resonance (NMR) spectroscopy.Alkyl-C (aliphatic compounds) and O-alkyl-C (carbohydrates) were the largest C fractions in soil and dissolved organic matter at Hubbard Brook. Alkyl-C ranged from 29–48% of soil C, 25–42% of soil solution C, and 22–42% of streamwater DOC. Carbohydrates comprised 32–49%, 36–43%, and 29–60% of C in soils, solutions, and streamwater, respectively. In both soils and soil solutions, the carbohydrate fraction decreased with increasing soil depth, while the aromaticity of organic matter increased with depth. There were no significant differences in the structural chemistry of SOM between clear-cut and uncut watersheds.The aromatic-C fractions in soil solutions at the clear-cut site ranged from 12–16%, approximately 40% greater than at the uncut site (8.5–11%). Thus, clear-cutting has resulted in the leaching of more highly decomposed organic matter, and depletion of more aliphatic compounds in the soluble organic pool. Because DOC fluxes are small compared to the SOM pool, large differences in soil solution chemistry do not substantially alter the overall composition of SOM. While the organic chemistry of stream DOC varied greatly among 3 sampling dates, there were no obvious clear-cutting effects. Thus, temporal variations in flowpaths and/or in-stream processes appear to be more important than disturbance in regulating the organic carbon chemistry of these streams.     

Micromonospora siamensis sp. nov., isolated from Thai peat swamp forest

Morphological and chemotaxonomic characterization of actinomycete strain TT2-4T isolated from peat swamp forest soil in Pattaloong Province, Thailand, clearly demonstrated that this strain belongs to the genus Micromonospora. 16S rDNA sequence analysis for the strain supported the assignment of the strain to the genus Micromonospora and the similarity value of sequences between this strain and the closely related species, Micromonospora mirobrigensis was 99.1%, and M. carbonacea and M. matsumotoense were 98.8%. The DNA-DNA hybridization result and some physiological and biochemical properties indicated that strain TT2-4T was distinguished from the phylogenetically closest relatives. Based on these genotypic and phenotypic data, strain TT2-4T merits a new species in the genus Micromonospora and the name Micromonospora siamensis sp. nov. is proposed for the strain. The type strain is strain TT2-4T (=JCM 12769T =PCU 266T =TISTR 1554T).     

Shorea balangeran and Dyera polyphylla (syn. Dyera lowii) as tropical peat swamp forest restoration transplant species: effects of mycorrhizae and level of disturbance

Tropical peat swamp forests (TPSF) are being rapidly deforested, leading to disturbed hydrology, wildfires and carbon loss. Cost-effective methods are needed to increase the scale of restoration activities. One method is to inoculate seedlings with their corresponding mycorrhizae species, thereby increasing performance during nursery cultivation, although the benefits post-transplantation are less well understood. This study considered two TPSF tree species, Shorea balangeran and Dyera polyphylla (syn. Dyera lowii), and their mycorrhiza; Scleroderma columnare (S. balangeran) and Glomus clarum and Gigaspora decipiens (D. polyphylla). The performance of non-inoculated and inoculated seedlings was compared following transplantation into five forest zones, representing a gradient from intact to degraded TPSF. In the degraded area, both inoculated seedling species supported higher colonization levels compared to non-inoculated seedlings. Both tree species showed high survival rates in all forest zones, and survival, growth and biomass production were not affected by mycorrhizal treatment. Both species grew faster and accumulated greater biomass in the more degraded forest zones. Nitrogen and phosphorus content reduced for both tree species in the more degraded forest zones, however, inoculated D. polyphylla seedlings had higher nutrient content across all forest zones, as did S. balangeran though less uniformly. Both these tree species are therefore suitable for reforesting degraded TPSF and mycorrhizal inoculation is recommended given a) inoculated seedlings in the degraded area permitted a higher mycorrhizal colonization level, and b) mycorrhizae increased nutrient uptake in the transplanted seedlings, although in this short-term study survival or growth improvement in the inoculated seedlings was not apparent.