Processes of organic carbon in mangrove ecosystems

In addition to carbon accumulation in plants, processes of organic carbon in mangrove ecosystems include origins of sediment organic carbon, carbon fluxes between mangroves and their adjacent systems (coastal waters and atmosphere), and cycling processes. Sediment organic carbon originates from suspending solids in coastal waters, mangrove plants and benthic algae. In mangroves with low organic carbon content in sediments, tidal seawater is the main origin of sediment organic carbon, while in mangroves with high sediment organic carbon contents, sediment organic carbon mainly originates from mangrove plants. Due to tidal flush, there is large material exchange between mangrove ecosystems and their adjacent coastal waters. In China, exports of organic carbon in litter falls and dissolved organic carbon from mangroves to their adjacent coastal waters have not been documented. Processes of mangrove litter falls, including production, decomposition, export and animal consumption, determine linkages among organic carbon among mangrove plants, secondary production and coastal ocean. Consumers especially benthic animals may influence organic carbon in mangrove ecosystems, because (1) their consumption rates are high, and their selective feeding on some food sources will change the relative quantities of export, bury and mineralization of organic carbon from different origins; (2) their consumption is much more than assimilation, resulting in the changes in sizes, forms and qualities of non-assimilated organic matters, and then the changes in availability of export, consumption or mineralization of organic carbon. Respiration and sulfate reduction are important mineralization processes of organic carbon in mangrove sediments. Mineralization rates of organic carbon in mangrove sediments are influenced by quantities, activities and particle sizes of organic matters, and other factors such as forest ages, root activities and animal burrowing activities. Researches on processes of mangrove organic carbon should be based on open systems, and ecological processes of organic carbon should be coupled with vegetation restoration.     

Processes of organic carbon in mangrove ecosystems

In addition to carbon accumulation in plants, processes of organic carbon in mangrove ecosystems include origins of sediment organic carbon, carbon fluxes between mangroves and their adjacent systems (coastal waters and atmosphere), and cycling processes. Sediment organic carbon originates from suspending solids in coastal waters, mangrove plants and benthic algae. In mangroves with low organic carbon content in sediments, tidal seawater is the main origin of sediment organic carbon, while in mangroves with high sediment organic carbon contents, sediment organic carbon mainly originates from mangrove plants. Due to tidal flush, there is large material exchange between mangrove ecosystems and their adjacent coastal waters. In China, exports of organic carbon in litter falls and dissolved organic carbon from mangroves to their adjacent coastal waters have not been documented. Processes of mangrove litter falls, including production, decomposition, export and animal consumption, determine linkages among organic carbon among mangrove plants, secondary production and coastal ocean. Consumers especially benthic animals may influence organic carbon in mangrove ecosystems, because (1) their consumption rates are high, and their selective feeding on some food sources will change the relative quantities of export, bury and mineralization of organic carbon from different origins; (2) their consumption is much more than assimilation, resulting in the changes in sizes, forms and qualities of non-assimilated organic matters, and then the changes in availability of export, consumption or mineralization of organic carbon. Respiration and sulfate reduction are important mineralization processes of organic carbon in mangrove sediments. Mineralization rates of organic carbon in mangrove sediments are influenced by quantities, activities and particle sizes of organic matters, and other factors such as forest ages, root activities and animal burrowing activities. Researches on processes of mangrove organic carbon should be based on open systems, and ecological processes of organic carbon should be coupled with vegetation restoration.     

能量标签技术及其在红树林生态系统能流研究中的应用

传统上认为红树林输出的有机质产生巨大的能流,支持了巨大的河口和近岸水域生态系统的次级生产。但能量标签技术的研究结果却显示红树林输出的有机质的作用并没有如此巨大。用红树碎屑难消化特性来解释此现象,此外数学模型模拟分析发现潮汐的稀释作用也可以解释这种现象。但这两者都不能解释,在其他初级生产者稀少时,红树材输出的有机质可以被大量利用的现象。在有红树林的河口和近海岸水域生态系统中,藻类等非红树初级生产者具有比红树植物更高的初级生产力,而且更容易被动物获得和消化。可以认为是藻类等巨大初级生产力的竞争作用导致红树初级生产在消费者组织中很难被发现,如此上面提到的难题就能得到很好的解决。此外能量标签技术检测出的是红树的初级生产在消费者组织中的相对比率,不是绝对数量值,从此角度看,能量标签技术的结果与传统观点不是矛盾而是互相补充的关系。由此推测红树的初级生产应该还是被消费者所利用,只是它们在消费者初级营养来源组成中占的比例并不大,但其绝对数量并不少。这与传统观点认为的红树的初级生产被大量利用,支撑了具有巨大的次级生产稍有不同。此外,能量标签技术在红树林生态系统中的适用性尚未检验;计算食物组成的数学工具不是很完善;实验设计上考虑的不够全面;对定量研究有一定的影响。     

Sediment biogeochemistry in an East African mangrove forest (Gazi Bay, Kenya)

The biogeochemistry of mangrove sediments was investigated in several mangrove forest communities in Gazi Bay, a coastal lagoon in Kenya, Africa. Carbon dioxide fluxes, sediment median grain sizes, sedimentary organic carbon, nitrogen and phosphorus contents and pore-water characteristics (ammonium, nitrate, sulfate and chloride) could be related to forest type. Mangrove sediments have pH values that range from 3.5 to 8.3 due to the limited buffer capacity of these sediments and intense acidifying processes such as aerobic degradation of organic matter, oxidation of reduced components, ammonium uptake by roots and root respiration. The mangrove sediments are nitrogen-rich compared to mangrove litter, as a result of microbial nitrogen retention, uptake and fixation, and import of nitrogen-rich material. It appears that mangrove sediments in Gazi Bay act as a nutrient and carbon sink rather than as a source for adjacent seagrass and reef ecosystems.     

Soil Respiration and Belowground Carbon Allocation in Mangrove Forests

Mangrove forests cover large areas of tropical and subtropical coastlines. They provide a wide range of ecosystem services that includes carbon storage in above- and below ground biomass and in soils. Carbon dioxide (CO₂) emissions from soil, or soil respiration is important in the global carbon budget and is sensitive to increasing global temperature. To understand the magnitude of mangrove soil respiration and the influence of forest structure and temperature on the variation in mangrove soil respiration I assessed soil respiration at eleven mangrove sites, ranging from latitude 27°N to 37°S. Mangrove soil respiration was similar to those observed for terrestrial forest soils. Soil respiration was correlated with leaf area index (LAI) and aboveground net primary production (litterfall), which should aid scaling up to regional and global estimates of soil respiration. Using a carbon balance model, total belowground carbon allocation (TBCA) per unit litterfall was similar in tall mangrove forests as observed in terrestrial forests, but in scrub mangrove forests TBCA per unit litter fall was greater than in terrestrial forests, suggesting mangroves allocate a large proportion of their fixed carbon below ground under unfavorable environmental conditions. The response of soil respiration to soil temperature was not a linear function of temperature. At temperatures below 26°C Q10 of mangrove soil respiration was 2.6, similar to that reported for terrestrial forest soils. However in scrub forests soil respiration declined with increasing soil temperature, largely because of reduced canopy cover and enhanced activity of photosynthetic benthic microbial communities.     

Increased litterfall in tropical forests boosts the transfer of soil CO2 to the atmosphere

Aboveground litter production in forests is likely to increase as a consequence of elevated atmospheric carbon dioxide (CO(2)) concentrations, rising temperatures, and shifting rainfall patterns. As litterfall represents a major flux of carbon from vegetation to soil, changes in litter inputs are likely to have wide-reaching consequences for soil carbon dynamics. Such disturbances to the carbon balance may be particularly important in the tropics because tropical forests store almost 30% of the global soil carbon, making them a critical component of the global carbon cycle; nevertheless, the effects of increasing aboveground litter production on belowground carbon dynamics are poorly understood. We used long-term, large-scale monthly litter removal and addition treatments in a lowland tropical forest to assess the consequences of increased litterfall on belowground CO(2) production. Over the second to the fifth year of treatments, litter addition increased soil respiration more than litter removal decreased it; soil respiration was on average 20% lower in the litter removal and 43% higher in the litter addition treatment compared to the controls but litter addition did not change microbial biomass. We predicted a 9% increase in soil respiration in the litter addition plots, based on the 20% decrease in the litter removal plots and an 11% reduction due to lower fine root biomass in the litter addition plots. The 43% measured increase in soil respiration was therefore 34% higher than predicted and it is possible that this 'extra' CO(2) was a result of priming effects, i.e. stimulation of the decomposition of older soil organic matter by the addition of fresh organic matter. Our results show that increases in aboveground litter production as a result of global change have the potential to cause considerable losses of soil carbon to the atmosphere in tropical forests.     

Biogeochemical effects of simulated sea level rise on carbon loss in an Everglades mangrove peat soil

Saltwater intrusion and inundation can affect soil microbial activity, which regulates the carbon (C) balance in mangroves and helps to determine if these coastal forests can keep pace with sea level rise (SLR). This study evaluated the effects of increased salinity (+15 ppt), increased inundation (?8 cm), and their combination, on soil organic C loss from a mangrove peat soil (Everglades, Florida, USA) under simulated tides. Soil respiration (CO₂ flux), methane (CH₄) flux, dissolved organic carbon (DOC) production, and porewater nutrient concentrations were quantified. Soil respiration was the major pathway of soil organic C loss (94–98%) and was approximately 90% higher in the control water level than the inundated treatment under elevated salinity. Respiration rate increased with water temperature, but depended upon salinity and tidal range. CH₄ flux was minimal, while porewater DOC increased with a concomitant, significant decline in soil bulk density under increased inundation. Porewater ammonium increased (73%) with inundation and soluble reactive phosphorus increased (32%) with salinity. Overall, the decline in soil organic C mineralization from combined saltwater intrusion and prolonged inundation was not significant, but results suggest SLR could increase this soil’s susceptibility to peat collapse and accelerate nutrient and DOC export to adjacent Florida Bay.     

Allometry, biomass, and productivity of mangrove forests: A review

We review 72 published articles to elucidate characteristics of biomass allocation and productivity of mangrove forests and also introduce recent progress on the study of mangrove allometry to solve the site- and species-specific problems. This includes the testing of a common allometric equation, which may be applicable to mangroves worldwide. The biomass of mangrove forests varies with age, dominant species, and locality. In primary mangrove forests, the above-ground biomass tends to be relatively low near the sea and increases inland. On a global scale, mangrove forests in the tropics have much higher above-ground biomass than those in temperate areas. Mangroves often accumulate large amounts of biomass in their roots, and the above-ground biomass to below-ground biomass ratio of mangrove forests is significantly low compared to that of upland forests (ANCOVA, P < 0.01). Several studies have reported on the growth increment of biomass and litter production in mangrove forests. We introduce some recent studies using the so-called “summation method” and investigate the trends in net primary production (NPP). For crown heights below 10 m, the above-ground NPP of mangrove forests is significantly higher (ANOVA, P < 0.01) than in those of tropical upland forests. The above-ground litter production is generally high in mangrove forests. Moreover, in many mangrove forests, the rate of soil respiration is low, possibly because of anaerobic soil conditions. These trends in biomass allocation, NPP, and soil respiration will result in high net ecosystem production, making mangrove forests highly efficient carbon sinks in the tropics.     

Carbon and nutrient exchange of mangrove forests with the coastal ocean

Mangrove forests exchange materials with the coastal ocean through tidal inundation. In this study, we aim to provide an overview of the published data of carbon (C) and nutrient exchange of mangrove forests with the coastal ocean at different spatial scales to assess whether the exchange is correlated with environmental parameters. We collected data on C (dissolved and particulate organic C; DOC and POC) and nutrient exchange (dissolved and particulate nitrogen, N and phosphorus, P) and examined the role of latitude, temperature, precipitation, geomorphological setting, hydrology, dominant mangrove species and forest area in explaining the variability of the exchange. We identified that there are a range of methodologies used to determine material exchange of mangroves with the coastal zone, each methodology providing data on the exchange at different spatial scales. This variability of approaches has limited our understanding of the role of mangroves in the coastal zone. Regardless, we found that mangrove forests export C and nutrients to the coastal zone in the form of litter and POC. We found that precipitation is a major factor influencing the export of C in the form of litter; sites with low annual precipitation and high mean annual temperatures export more C as litter than sites with high precipitation and low temperature. Furthermore, export of POC is higher in zones with low mean annual minimum temperature. Identification of broad-scale trends in DOC and dissolved nutrients was more difficult, as the analysis was limited by scarcity of suitable studies and high variability in experimental approaches. However, tidal amplitude and the concentration of nutrients in the floodwater appears to be important in determining nutrient exchange. The strongest conclusion from our analysis is that mangrove forests are in general sources of C and nutrients in the form of litter and POC and that they are most likely to be exporting C subsidies in dry regions.     

Plant-animal interactions and the structure and function of mangrove forest ecosystems*

Trophic interactions involving plants and animals in tropical mangrove forests have important controlling influences on several population, community and ecosystem-level processes. Insect herbivores remove up to 35% of leaf area from some mangrove tree species and can cause the death of seedlings. Leaf chemistry and toughness and soil nutrient status all appear to be important in explaining the between- and among-species variance in leaf damage. Insects also attack and damage, mainly by boring, a large proportion of mangrove seeds. Shadehouse experiments have shown that such post-dispersal predation can have a significant effect on seedling survival, growth and biomass allocation to leaves, stems and roots. Sesarmid crabs are also responsible for severe post-dispersal seed predation. In field trials, crabs consumed more than 70% of the seeds of five tree species. For four of these five species there was an inverse relationship between seed predation rate and the dominance of conspecific adult trees, while the within-site distribution pattern of one tree species appears to be partially controlled by crabs. The same crab species also consume 30–80% (depending on forest type and intertidal elevation), of the annual litter fall in mangrove forests and, thus, have an important role in controlling the rate of remineralization of detritus within forests and the export of particulate matter from the forests to other nearshore habitats. The other major component of litter in the forests is wood, which is broken down relatively rapidly by teredinid molluscs (shipworms). More than 90% of the weight loss from decomposing trunks of Rhizophora species during the first four years of decay is through ingestion by teredinids. The annual turnover of dead wood mass in Rhizophora forests is equivalent to that of the processing of leaf detritus by crabs. Because of the relatively low species richness of trees and consumers in tropical mangrove forests, they are likely to serve as productive sites for further investigations of the influence of plant-animal interactions on the dynamics of tropical forests.     

Factors regulating carbon sinks in mangrove ecosystems

Mangroves are recognized as one of the richest carbon storage systems. However, the factors regulating carbon sinks in mangrove ecosystems are still unclear, particularly in the subtropical mangroves. The biomass, production, litterfall, detrital export and decomposition of the dominant mangrove vegetation in subtropical (Kandelia obovata) and tropical (Avicennia marina) Taiwan were quantified from October 2011 to July 2014 to construct the carbon budgets. Despite the different tree species, a principal component analysis revealed the site or environmental conditions had a greater influence than the tree species on the carbon processes. For both species, the net production (NP) rates ranged from 10.86 to 27.64 Mg C ha^?1 year^?1 and were higher than the global average rate due to the high tree density. While most of the litterfall remained on the ground, a high percentage (72%–91%) of the ground litter decomposed within 1 year and fluxed out of the mangroves. However, human activities might cause a carbon flux into the mangroves and a lower NP rate. The rates of the organic carbon export and soil heterotrophic respiration were greater than the global mean values and those at other locations. Only a small percentage (3%–12%) of the NP was stored in the sediment. The carbon burial rates were much lower than the global average rate due to their faster decomposition, indicating that decomposition played a critical role in determining the burial rate in the sediment. The summation of the organic and inorganic carbon fluxes and soil heterotrophic respiration well exceeded the amount of litter decomposition, indicating an additional source of organic carbon that was unaccounted for by decomposition in the sediment. Sediment‐stable isotope analyses further suggest that the trapping of organic matter from upstream rivers or adjacent waters contributed more to the mangrove carbon sinks than the actual production of the mangrove trees.     

重金属污染对珠江口红树林表层沉积物碳含量的影响

湿地沉积物是红树林生态系统中重要的组成部分,其总有机碳储量的变化对红树林生态系统的固碳能力有着重要影响。现有对红树林湿地重金属的研究多集中于污染评价,鲜有涉及重金属含量对沉积物总有机碳（TOC）含量影响的研究。于2018-2019年期间4次前往珠江口典型红树林湿地,采集了0-30 cm表层土壤沉积物的样品,并测定其重金属含量和TOC含量,以探讨重金属含量变化对TOC的影响。结果表明,与广东地区的背景值相比,研究区沉积物重金属含量超标较为严重,重金属来源应是人类活动;沉积物的重金属含量能够显著影响TOC含量（P<0.01,R²=0.39）,间接对红树林湿地的固碳能力、甚至全球变暖产生一定影响;Cd、As、Zn含量高的沉积物环境有利于TOC的积累,Cu、Cr、Ni、Hg含量低的沉积物环境则不利于TOC的积累。沉积物的重金属对TOC的影响的机制是非常复杂的,它们可以通过影响土壤结构、土壤化学组分、土壤内微生物、上部植被群落的生长以及凋落物归还等一系列过程,导致沉积物TOC和固碳能力的变化。     

Microbial growth and detritus transformations during decomposition of leaf litter in a stream

SUMMARY. 1. Despite the widely accepted importance of bacteria and fungi in degrading detritus in aquatic ecosystems there is still very little quantitative information on the abundance and dynamics of these microorganisms. Using epifluorescent microscopy, we measured the biomass of bacteria and fungi during decomposition of three types of leaf detritus. Bacterial production was determined from the rate of incorporation of ³H-thymidine into DNA.
2. The transformation of leaf carbon into dissolved organic carbon and fine particulate organic carbon was followed in order to compare the amounts of leaf material that were converted into these 'end-products' of decomposition versus the amount converted into microbial biomass.
3. The amount of microbial carbon in the leaf-detritus complex never exceeded 5.2% of the total carbon, and fungal biomass was always much greater than bacterial biomass. Despite the greater standing stock of fungi, the rapid turnover of bacteria (doubling about once per day) implies that their role in degrading leaf litter or as a food source for detritivores might be as great as for fungi.
4. Removal of microbial biomass from leaf litter may occur as release of fungal spores and consumption or shedding of bacterial biomass. Fungal spores can be a significant part of the fine particulate organic carbon released from leaf detritus and potentially represent an important food resource for filter-feeding organisms.     

Effect of soil microorganisms and labile C availability on soil respiration in response to litter inputs in forest ecosystems: A meta‐analysis

Litter inputs can influence soil respiration directly through labile C availability and, indirectly, through the activity of soil microorganisms and modifications in soil microclimate; however, their relative contributions and the magnitude of any effect remain poorly understood. We synthesized 66 recently published papers on forest ecosystems using a meta‐analysis approach to investigate the effect of litter inputs on soil respiration and the underlying mechanisms involved. Our results showed that litter inputs had a strong positive impact on soil respiration, labile C availability, and the abundance of soil microorganisms, with less of an impact related to soil moisture and temperature. Overall, soil respiration was increased by 36% and 55%, respectively, in response to natural and doubled litter inputs. The increase in soil respiration induced by litter inputs showed a tendency for coniferous forests (50.7%)> broad‐leaved forests (41.3%)> mixed forests (31.9%). This stimulation effect also depended on stand age with 30‐ to 100‐year‐old forests (53.3%) and ≥100‐year‐old forests (50.2%) both 1.5 times larger than ≤30‐year‐old forests (34.5%). Soil microbial biomass carbon and soil dissolved organic carbon increased by 21.0%‐33.6% and 60.3%‐87.7%, respectively, in response to natural and doubled litter inputs, while soil respiration increased linearly with corresponding increases in soil microbial biomass carbon and soil dissolved organic carbon. Natural and doubled litter inputs increased the total phospholipid fatty acid (PLFA) content by 6.6% and 19.7%, respectively, but decreased the fungal/bacterial PLFA ratio by 26.9% and 18.7%, respectively. Soil respiration also increased linearly with increases in total PLFA and decreased linearly with decreases in the fungal/bacterial PLFA ratio. The contribution of litter inputs to an increase in soil respiration showed a trend of total PLFA > fungal/bacterial PLFA ratio > soil dissolved organic carbon > soil microbial biomass carbon. Therefore, in addition to forest type and stand age, labile C availability and soil microorganisms are also important factors that influence soil respiration in response to litter inputs, with soil microorganisms being more important than labile C availability.     

A simulation model of organic matter and nutrient accumulation in mangrove wetland soils

The distribution and accumulation of organic matter, nitrogen (N) and phosphorus (P) in mangrove soils at four sites along the Shark River estuary of south Florida were investigated with empirical measures and a process-based model. The mangrove nutrient model (NUMAN) was developed from the SEMIDEC marsh organic matter model and parameterized with data from mangrove wetlands. The soil characteristics in the four mangrove sites varied greatly in both concentrations and profiles of soil carbon, N and P. Organic matter decreased from 82% in the upstream locations to 30% in the marine sites. Comparisons of simulated and observed results demonstrated that landscape gradients of soil characteristics along the estuary can be adequately modeled by accounting for plant production, litter decomposition and export, and allochthonous input of mineral sediments. Model sensitivity analyses suggest that root production has a more significant effect on soil composition than litter fall. Model simulations showed that the greatest change in organic matter, N, and P occurred from the soil surface to 5 cm depth. The rapid decomposition of labile organic matter was responsible for this decrease in organic matter. Simulated N mineralization rates decreased quickly with depth, which corresponded with the decrease of labile organic matter. The increase in organic matter content and decrease in soil bulk density from mangrove sites at downstream locations compared to those at upstream locations was controlled mainly by variation in allochthonous inputs of mineral matter at the mouth of the estuary, along with gradients in mangrove root production. Research on allochthonouns sediment input and in situ root production of mangroves is limited compared to their significance to understanding nutrient biogeochemistry of these wetlands. More accurate simulations of temporal patterns of nutrient characteristics with depth will depend on including the effects of disturbance such as hurricanes on sediment redistribution and biomass production.     

The influence of crabs on litter processing in high intertidal mangrove forests in tropical Australia

Summary Measurements of litter fall and litter removal by crabs, in conjunction with estimates of litter decay by microbes and tidal export of litter from three high-intertidal mangrove forests were made during a year-long study in tropical northeastern Australia. In forests dominated by Ceriops tagal and Bruguiera exaristata, litter standing stocks remained low on the forest floor (mean 6 g·m^-2), although litter fall was high; 822 and 1022 g·m^-2·y^-1, respectively. Sesarmid crabs removed 580 (Ceriops) and 803 (Bruguiera) g·m^-2·y^-1, or 71 and 79%, of the total annual litter fall from the forest floor. Relative to the rate of litter removal by crabs, microbial turnover of whole, unshredded litter was insignificant, accounting for <1% of annual litter fall. Export of litter by tides was estimated to remove 194 (Ceriops) and 252 (Bruguiera) g·m^-2·y^-1 or 24 and 25% of annual litter fall. In a forest dominated by Avicenniamarina, in which an ocypodid crab was more abundant than sesarmids, litter standing stocks were higher (mean 84 g·m^-2) and crabs removed less litter; 173 g·m^-2·y^-1 or 33% of the annual litter fall of 519 g·m^-2·y^-1. Microbial turnover of intact litter was more important in the Avicennia forest (168 g·m^-2·y^-1 or 32% of annual litter fall), and tides exported 107 g·m^-2·y^-1 or 21% of litter production. In areas where sesarmid crabs were absent or rare in Ceriops forests, there were significantly higher standing stocks of litter and slower rates of leaf removal. Taking into account the probable assimilation efficiencies of sesarmid crabs feeding on mangrove leaves, we estimate that in Ceriops and Bruguiera forests leaf processing by crabs turns litter over at >75 times the rate of microbial decay alone, thus facilitating the high sediment bacterial productivity in these forests. The importance of litter processing by crabs increases with height in the intertidal in tropical Australia, in contrast to New World mangrove forests, where the reverse is true.Contribution No. 445 from the Australian Institute of Marine Science     

Fate of leaf litter in restored Kandelia obovata (S. L.) mangrove forests with different ages in Jiulong River Estuary,China

Ecological processing of leaf litter plays important roles in carbon dynamics of mangrove forests. Fate of leaf litter, that is, removal by crabs, microbial decomposition, and tidal export was quantified in two restored Kandelia obovata forests with ages of 24 years and 48 years, respectively, from December 2009 to November 2010. Crab abundance was also investigated to test the role of crabs in leaf litter processing. Daily leaf litter production was 1.064 ± 0.108 g C m^?2 day^?1 at the 24‐year forest and was 0.689 ± 0.040 g C m^?2 day^?1 at the 48‐year forest. Annual mean removal of leaf litter by crabs was lower at the 24‐year forest than at the 48‐year forest (0.177 ± 0.046 g C m^?2 day^?1 vs. 0.220 ± 0.050 g C m^?2 day^?1), due to a higher crab abundance at the older forest. Microbial decomposition and change in standing stock of leaf litter on the forest floor made a negligible contribution to the annual leaf litter production. Tidal exports of leaf litter were estimated as 0.875 ± 0.090 g C m^?2 day^?1 and 0.458 ± 0.086 g C m^?2 day^?1 at the 24‐year and 48‐year forests, respectively, accounting for 82.2% and 66.5% of their daily leaf litter production. Turnover rate of leaf litter was higher at the younger forest (1.7 ± 0.4 day^?1) than the older forest (1.2 ± 0.3 day^?1). Removal of leaf litter by crabs was higher in warm months while tidal export of leaf litter showed a much less apparent seasonal pattern. Spatial variations of crab removal and tidal export of leaf litter with forest zones were observed within each forest, while microbial decomposition of leaf litter was comparable among the different zones. These indicated that the ecosystem functions of restored mangrove forest could not reach a level equivalent to those of a mature forest even 24 years after restoration.     

Mangrove outwelling: a review

The export of detritus and faunal biomass from mangroves has long been considered as an important support for offshore biological production and has been widely used as an argument for mangrove conservation. This functional role of the mangroves, like many other paradigms in mangrove ecology, has seldom been put to rigorous test since the hypothesis was postulated about 25 years ago. Past studies on which the hypothesis was based were mostly carried out in mangrove or other wetland environments, little is known about the fate and effects of outwelled detritus on oceanic, offshore, communities. Mass balance studies carried out in the last 15 years tend to suggest that export is common from tidal mangroves, the direction of flow depends, however, on the identity of the chemical species in question. Pore water and groundwater flow can affect tidal material exchange but are poorly studied. Generally, tracer methods using stable isotope ratios or other signatures have suggested that outwelling may be much less significant than expected. Further, most past studies have focussed on particulate matter while it is increasingly apparent that dissolved organics may play a more important role in matter exchange between offshore and mangrove communities. There is also evidence that benthic biomass and richness may not bear any positive or significant relationship with detritus availability.     

Spatial and temporal variation of microbial respiration rates in a blackwater stream

1. The extent of spatial and temporal variation of microbial respiration was determined in a first-order, sand-bottomed, blackwater stream on the coastal plain of south-eastern Virginia, U.S.A.
2. Annual mean respiration rates (as g O₂ m^–3 h^–1) differed significantly among substrata: leaf litter, 12.9; woody debris, 2.4; surface sediment, 0.8; hyporheic sediment, 0.4; water column, 0.003. Rates associated with wood were higher than those with leaves when expressed per unit surface area.
3. Highest respiration rates on leaves, wood and in the water column occurred during the summer, whereas rates in the sediments were greatest during the late autumn and winter. Water temperature, as well as particulate organic matter and nitrogen content of the substrata, was correlated positively with respiration rates.
4. A stepwise multiple regression showed that temperature and nitrogen content together explained 88% of the variation in respiration rates of leaves and wood. In contrast, particulate organic matter content and nitrogen content explained 89–90% of the variation in respiration in the sediments. Although water temperature was a significant factor in the sediment multiple regressions, its addition as an independent variable improved the regression models only slightly.
5. Annual mean respiration in the stream channel, based on the proportional amount of respiration occurring associated with each type of substratum during each month, was 1.1 kg O₂ m^–2 yr^–1. Seventy per cent of respiration in the stream occurred in the hyporheic zone, 8–13% occurred in the surface sediment, leaf litter or woody debris, and < 1% occurred in the water column. Approximately 16% of total detritus, or 40% of non-woody detritus, stored in the stream during the year was lost to microbial respiration.     

Benthic bacterial and fungal productivity and carbon turnover in a freshwater marsh

Heterotrophic bacteria and fungi are widely recognized as crucial mediators of carbon, nutrient, and energy flow in ecosystems, yet information on their total annual production in benthic habitats is lacking. To assess the significance of annual microbial production in a structurally complex system, we measured production rates of bacteria and fungi over an annual cycle in four aerobic habitats of a littoral freshwater marsh. Production rates of fungi in plant litter were substantial (0.2 to 2.4 mg C g(-1) C) but were clearly outweighed by those of bacteria (2.6 to 18.8 mg C g(-1) C) throughout the year. This indicates that bacteria represent the most actively growing microorganisms on marsh plant litter in submerged conditions, a finding that contrasts strikingly with results from both standing dead shoots of marsh plants and submerged plant litter decaying in streams. Concomitant measurements of microbial respiration (1.5 to 15.3 mg C-CO2 g(-1) of plant litter C day(-1)) point to high microbial growth efficiencies on the plant litter, averaging 45.5%. The submerged plant litter layer together with the thin aerobic sediment layer underneath (average depth of 5 mm) contributed the bulk of microbial production per square meter of marsh surface (99%), whereas bacterial production in the marsh water column and epiphytic biofilms was negligible. The magnitude of the combined production in these compartments (approximately 1,490 g C m(-2) year(-1)) highlights the importance of carbon flows through microbial biomass, to the extent that even massive primary productivity of the marsh plants (603 g C m(-2) year(-1)) and subsidiary carbon sources (approximately 330 g C m(-2) year(-1)) were insufficient to meet the microbial carbon demand. These findings suggest that littoral freshwater marshes are genuine hot spots of aerobic microbial carbon transformations, which may act as net organic carbon importers from adjacent systems and, in turn, emit large amounts of CO2 (here, approximately 870 g C m(-2) year(-1)) into the atmosphere.