Mangrove tannins in aquatic ecosystems: Their fate and possible influence on dissolved organic carbon and nitrogen cycling

We describe the fate of mangrove leaf tannins in aquatic ecosystems and their possible influence on dissolved organic nitrogen (DON) cycling. Tannins were extracted and purified from senescent yellow leaves of the red mangrove (Rhizophora mangle) and used for a series of model experiments to investigate their physical and chemical reactivity in natural environments. Physical processes investigated included aggregation, adsorption to organic matter-rich sediments, and co-aggregation with DON in natural waters. Chemical reactions included structural change, which was determined by excitation–emission matrix fluorescence spectra, and the release of proteins from tannin–protein complexes under solar-simulated light exposure. A large portion of tannins can be physically eliminated from aquatic environments by precipitation in saline water and also by binding to sediments. A portion of DON in natural water can coprecipitate with tannins, indicating that mangrove swamps can influence DON cycling in estuarine environments. The chemical reactivity of tannins in natural waters was also very high, with a half-life of less than 1 d. Proteins were released gradually from tannin–protein complexes incubated under light conditions but not under dark conditions, indicating a potentially buffering role of tannin– protein complexes on DON recycling in mangrove estuaries. Although tannins are not detected at a significant level in natural waters, they play an important ecological role by preserving nitrogen and buffering its cycling in estuarine ecosystems through the prevention of rapid DON export/loss from mangrove fringe areas and/or from rapid microbial mineralization.

Quantitative and Qualitative Aspects of Dissolved Organic Carbon Leached from Senescent Plants in an Oligotrophic Wetland

We conducted a series of experiments whereby dissolved organic matter (DOM) was leached from various wetland and estuarine plants, namely sawgrass (Cladium jamaicense), spikerush (Eleocharis cellulosa), red mangrove (Rhizophora mangle), cattail (Typha domingensis), periphyton (dry and wet mat), and a seagrass (turtle grass; Thalassia testudinum). All are abundant in the Florida Coastal Everglades (FCE) except for cattail, but this species has a potential to proliferate in this environment. Senescent plant samples were immersed into ultrapure water with and without addition of 0.1% NaN3 (w/ and w/o NaN3, respectively) for 36 days. We replaced the water every 3 days. The amount of dissolved organic carbon (DOC), sugars, and phenols in the leachates were analyzed. The contribution of plant leachates to the ultrafiltered high molecular weight fraction of DOM (>1 kDa; UDOM) in natural waters in the FCE was also investigated. UDOM in plant leachates was obtained by tangential flow ultrafiltration and its carbon and phenolic compound compositions were analyzed using solid state 13C cross-polarization magic angle spinning nuclear magnetic resonance (13C CPMAS NMR) spectroscopy and thermochemolysis in the presence of tetramethylammonium hydroxide (TMAH thermochemolysis), respectively. The maximum yield of DOC leached from plants over the 36-day incubations ranged from 13.0 to 55.2 g C kg−1 dry weight. This amount was lower in w/o NaN3 treatments (more DOC was consumed by microbes than produced) except for periphyton. During the first 2 weeks of the 5 week incubation period, 60–85% of the total amount of DOC was leached, and exponential decay models fit the leaching rates except for periphyton w/o NaN3. Leached DOC (w/ NaN3) contained different concentrations of sugars and phenols depending on the plant types (1.09–7.22 and 0.38–12.4 g C kg−1 dry weight, respectively), and those biomolecules comprised 8–34% and 4–28% of the total DOC, respectively. This result shows that polyphenols that readily leach from senescent plants can be an important source of chromophoric DOM (CDOM) in wetland environments. The O-alkyl C was found to be the major C form (55±9%) of UDOM in plant leachates as determined by 13C CPMAS NMR. The relative abundance of alkyl C and carbonyl C was consistently lower in plant-leached UDOM than that in natural water UDOM in the FCE, which suggests that these constituents increase in relative abundance during diagenetic processing. TMAH thermochemolysis analysis revealed that the phenolic composition was different among the UDOM leached from different plants, and was expected to serve as a source indicator of UDOM in natural water. Polyphenols are, however, very reactive and photosensitive in aquatic environments, and thus may loose their plant-specific molecular characteristics shortly. Our study suggests that variations in vegetative cover across a wetland landscape will affect the quantity and quality of DOM leached into the water, and such differences in DOM characteristics may affect other biogeochemical processes.

Changes in mass and nutrient content of wood during decomposition in a south Florida mangrove forest

1. Large pools of dead wood in mangrove forests following disturbances such as hurricanes may influence nutrient fluxes. We hypothesized that decomposition of wood of mangroves from Florida, USA (Avicennia germinans, Laguncularia racemosa and Rhizophora mangle), and the consequent nutrient dynamics, would depend on species, location in the forest relative to freshwater and marine influences and whether the wood was standing, lying on the sediment surface or buried. 2. Wood disks (8–10 cm diameter, 1 cm thick) from each species were set to decompose at sites along the Shark River, either buried in the sediment, on the soil surface or in the air (above both the soil surface and high tide elevation). 3. A simple exponential model described the decay of wood in the air, and neither species nor site had any effect on the decay coefficient during the first 13 months of decomposition. 4. Over 28 months of decomposition, buried and surface disks decomposed following a two-component model, with labile and refractory components. Avicennia germinans had the largest labile component (18 ± 2% of dry weight), while Laguncularia racemosa had the lowest (10 ± 2%). Labile components decayed at rates of 0.37–23.71% month−1, while refractory components decayed at rates of 0.001–0.033% month−1. Disks decomposing on the soil surface had higher decay rates than buried disks, but both were higher than disks in the air. All species had similar decay rates of the labile and refractory components, but A. germinans exhibited faster overall decay because of a higher proportion of labile components. 5. Nitrogen content generally increased in buried and surface disks, but there was little change in N content of disks in the air over the 2-year study. Between 17% and 68% of total phosphorus in wood leached out during the first 2 months of decomposition, with buried disks having the greater losses, P remaining constant or increasing slightly thereafter. 6. Newly deposited wood from living trees was a short-term source of N for the ecosystem but, by the end of 2 years, had become a net sink. Wood, however, remained a source of P for the ecosystem. 7. As in other forested ecosystems, coarse woody debris can have a significant impact on carbon and nutrient dynamics in mangrove forests. The prevalence of disturbances, such as hurricanes, that can deposit large amounts of wood on the forest floor accentuates the importance of downed wood in these forests.

Leaf Gas Exchange and Nutrient Use Efficiency Help Explain the Distribution of Two Neotropical Mangroves under Contrasting Flooding and Salinity

Rhizophora mangle and Laguncularia racemosa cooccur along many intertidal floodplains in the Neotropics. Their patterns of dominance shift along various gradients, coincident with salinity, soil fertility, and tidal flooding. We used leaf gas exchange metrics to investigate the strategies of these two species in mixed culture to simulate competition under different salinity concentrations and hydroperiods. Semidiurnal tidal and permanent flooding hydroperiods at two constant salinity regimes (10 g L−1 and 40 g L−1) were simulated over 10 months. Assimilation ( ), stomatal conductance ( ), intercellular CO2 concentration ( ), instantaneous photosynthetic water use efficiency (PWUE), and photosynthetic nitrogen use efficiency (PNUE) were determined at the leaf level for both species over two time periods. Rhizophora mangle had significantly higher PWUE than did L. racemosa seedlings at low salinities; however, L. racemosa had higher PNUE and and, accordingly, had greater intercellular CO2 (calculated) during measurements. Both species maintained similar capacities for A at 10 and 40 g L−1 salinity and during both permanent and tidal hydroperiod treatments. Hydroperiod alone had no detectable effect on leaf gas exchange. However, PWUE increased and PNUE decreased for both species at 40 g L−1 salinity compared to 10 g L−1. At 40 g L−1 salinity, PNUE was higher for L. racemosa than R. mangle with tidal flooding. These treatments indicated that salinity influences gas exchange efficiency, might affect how gases are apportioned intercellularly, and accentuates different strategies for distributing leaf nitrogen to photosynthesis for these two species while growing competitively.

Pollen distribution of sediment core MD96-2048 from the Western Indian Ocean

Glacial-interglacial fluctuations in the vegetation of South Africa might elucidate the climate system at the edge of the tropics between the Indian and Atlantic Oceans. However, vegetation records covering a full glacial cycle have only been published from the eastern South Atlantic. We present a pollen record of the marine core MD96-2048 retrieved by the Marion Dufresne from the Indian Ocean ~120 km south of the Limpopo River mouth. The sedimentation at the site is slow and continuous. The upper 6 m (spanning the past 342 Ka) have been analysed for pollen and spores at millennial resolution. The terrestrial pollen assemblages indicate that during interglacials, the vegetation of eastern South Africa and southern Mozambique largely consisted of evergreen and deciduous forests. During glacials open mountainous scrubland dominated. Montane forest with Podocarpus extended during humid periods was favoured by strong local insolation. Correlation with the sea surface temperature record of the same core indicates that the extension of mountainous scrubland primarily depends on sea surface temperatures of the Agulhas Current. Our record corroborates terrestrial evidence of the extension of open mountainous scrubland (including fynbos-like species of the high-altitude Grassland biome) for the last glacial as well as for other glacial periods of the past 300 Ka.

Palaeoenvironment records of six sediment cores collected in the Gulf of Carpentaria

The Gulf of Carpentaria is an epicontinental sea (maximum depth 70 m) between Australia and New Guinea, bordered to the east by Torres Strait (currently 12 m deep) and to the west by the Arafura Sill (53 m below present sea level). Throughout the Quaternary, during times of low sea-level, the Gulf was separated from the open waters of the Indian and Pacific Oceans, forming Lake Carpentaria, an isolation basin, perched above contemporaneous sea-level with outlet channels to the Arafura Sea. A preliminary interpretation is presented of the palaeoenvironments recorded in six sediment cores collected by the IMAGES program in the Gulf of Carpentaria. The longest core (approx. 15 m) spans the past 130 ka and includes a record of sea-level/lake-level changes, with particular complexity between 80 and 40 ka when sea-level repeatedly breached and withdrew from Gulf/Lake Carpentaria. Evidence from biotic remains (foraminifers, ostracods, pollen), sedimentology and geochemistry clearly identifies a final marine transgression at about 9.7 ka (radiocarbon years). Before this transgression, Lake Carpentaria was surrounded by grassland, was near full, and may have had a surface area approaching 600 km-300 km and a depth of about 15 m. The earlier rise in sea-level which accompanied the Marine Isotopic Stage 6/5 transgression at about 130 ka is constrained by sedimentological and biotic evidence and dated by optical- and thermoluminescence and amino acid racemisation methods.

Pollen and spore counts of sediment core GeoB12624

In tropical eastern Africa, vegetation distribution is largely controlled by regional hydrology, which has varied over the past 20 000 years. Therefore, accurate reconstructions of past vegetation and hydrological changes are crucial for a better understanding of climate variability in the tropical southeastern African region. We present high-resolution pollen records from a marine sediment core recovered offshore of the Rufiji River delta. Our data document significant shifts in pollen assemblages during the last deglaciation, identifying, through changes in both upland and lowland vegetation, specific responses of plant communities to atmospheric (precipitation) and coastal (coastal dynamics and sea-level changes) alterations. Specifically, arid conditions reflected by a maximum pollen representation of dry and open vegetation occurred during the Northern Hemisphere cold Heinrich event 1 (H1), suggesting that the expansion of drier upland vegetation was synchronous with cold Northern Hemisphere conditions. This arid period is followed by an interval in which forest and humid woodlands expanded, indicating a hydrologic shift towards more humid conditions. Droughts during H1 and the shift to humid conditions around 14.8 kyr BP in the uplands are consistent with latitudinal shifts of the intertropical convergence zone (ITCZ) driven by high-latitude Northern Hemisphere climatic fluctuations. Additionally, our results show that the lowland vegetation, consisting of well-developed salt marshes and mangroves in a successional pattern typical for vegetation occurring in intertidal habitats, has responded mainly to local coastal dynamics related to marine inundation frequencies and soil salinity in the Rufiji Delta as well as to the local moisture availability. Lowland vegetation shows a substantial expansion of mangrove trees after ~ 14.8 kyr BP, suggesting an increased moisture availability and river runoff in the coastal area. The results of this study highlight the decoupled climatic and environmental processes to which the vegetation in the uplands and the Rufiji Delta has responded during the last deglaciation.

Age model and pollen analysis from site GIK16867 in the northern Angola Basin

Sporomorphs and dinoflagellate cysts from site GIK16867 in the northern Angola Basin record the vegetation history of the West African forest during the last 700 ka in relation to changes in salinity and productivity of the eastern Gulf of Guinea. During most cool and cold periods, the Afromontane forest, rather than the open grass-rich dry forest, expanded to lower altitudes partly replacing the lowland rain forest of the borderlands east of the Gulf of Guinea. Except in Stage 3, when oceanic productivity was high during a period of decreased atmospheric circulation, high oceanic productivity is correlated to strong winds. The response of marine productivity in the course of a climatic cycle, however, is earlier than that of wind vigour and makes wind-stress-induced oceanic upwelling in the area less likely. Monsoon variation is well illustrated by the pollen record of increased lowland rain forest that is paired to the dinoflagellate cyst record of decreased salinity forced by increased precipitation and run-off.

Pollen flux off Cape Blanc

Fluxes of airborne freshwater diatoms (FD), phytoliths (PH), and pollen grains (PO) collected with sediment traps off Cape Blanc, northwest Africa, from 1988 till 1991 are presented. Both continental rainfall variations and wind mean strength and direction play a key role in the temporal fluctuations of the fluxes of eolian traces in the pelagic realm. Drier conditions in Northern Africa in 1987 could have preceded the high lithogenic input and moderate FD flux in 1988. The PH peak in summer 1988 was probably caused by increased wind velocity. Wetter rainy seasons of 1988/89 might have promoted a significant pollen production in summer 1989, and FD in late 1989 and early 1990, as well as contributed to the reduction of the lithogenic flux in 1989/90. Decreased fluxes of FD, PH and PO, and higher contribution of the 6-11 µm lithogenic fraction in 1991 would mainly reflect minor intensity and decreased amount of continental trade winds. Air-mass backward trajectories confirm that the Saharan Air Layer is predominantly involved in the spring/summer transport. Trade winds play a decisive role in the fall/winter months, but also contribute to the transport during late spring/summer. Origin of wind trajectories does not support a direct relationship between transporting wind-layers and material source areas in Northern Africa. High winter fluxes of eolian tracers and high amount of trade winds with continental origin in summer warn against a simplistic interpretation of the seasonal eolian signal preserved in the sediments off Cape Blanc, and the wind layer involved in its transport.

Pollen analysis of sediment core GeoB1023-5

A high resolution marine pollen record from site GeoB1023, west of the northern Namib desert provides data on vegetation and climate change for the last 21 ka at an average resolution of 185 y. Pollen and spores are mainly delivered to the site by the Cunene river and by surface and mid-tropospheric wind systems. The main pollen source areas are located between 13°S and 21°S, which includes the northern Namib desert and semi-desert, the Angola-northern Namibian highland, and the north-western Kalahari. The pollen spectra reflect environmental changes in the region. The last glacial maximum (LGM) was characterised by colder and more arid conditions than at present, when a vegetation with temperate elements such as Asteroideae, Ericaceae, and Restionaceae grew north of 21°S. At 17.5 ka cal. B.P., an amelioration both in temperature and humidity terminated the LGM but, in the northern Kalahari, mean annual rainfall in the interval 17.5-14.4 ka cal. B.P. was probably 100-150 mm lower than at present (400-500 mm/y). The Late-glacial to early Holocene transition includes two arid periods, i.e. 14.4-12.5 and 10.9-9.3 ka cal. B.P. The last part of the former period may be correlated with the Younger Dryas. The warmest and most humid period in the Holocene occurred between 6.3 and 4.8 ka cal. B.P. During the last 2000 years, human impact, as reflected by indications of deforestation, enhanced burning and overgrazing, progressively intensified.

Pollen record of sediment core GeoB1008-3 from the Congo fan

Palynological investigation of the marine core, GeoB1008-3, from near the mouth of the Congo river (6°35.6'S/10°19.1'E), provides information about the changes in vegetation and climate in West Equatorial Africa during the last 190 ka. The pollen diagram is divided into zones 1-6 which are considered to correspond in time with the marine isotope stages 1-6. Oscillations in temperature and moisture are indicated during the cold stage 6. During stage 5, two cooler periods (5d and 5b) can be shown with an expansion of Podocarpus forests to lower elevations on the expense of lowland rain forest. Extended mangrove swamps existed along the coast in times of high sea level (stages 5 and 1).

Pollen analysis of sediment core GIK16415-2 in the eastern tropical Atlantic

Palynological data of the marine core M 16415-2 show latitudinal shifts of the northern fringe of the tropical rain forest in north-west Africa during the last 700 ka. Savanna and dry open forest expanded southwards and tropical rain forest expanded northwards during dry and humid periods, respectively. Until 220 ka B.P., the tropical rain forest probably kept its zonal character in West Africa during glacials and interglacials. It is only during the last two glacial periods that the rain forest possibly fragmented into refugia. Throughout the Brunhes chron, pollen and spore transport was mainly by trade winds.

Evaluating, predicting and mapping belowground carbon stores in Kenyan mangroves.

Despite covering only approximately 138,000 km2, mangroves are globally important carbon sinks with carbon density values 3 to 4 times that of terrestrial forests. A key challenge in evaluating the carbon benefits from mangrove forest conservation is the lack of rigorous spatially resolved estimates of mangrove sediment carbon stocks; most mangrove carbon is stored belowground. Previous work has focused on detailed estimations of carbon stores over relatively small areas, which has obvious limitations in terms of generality and scope of application. Most studies have focused only on quantifying the top 1m of belowground carbon (BGC). Carbon stored at depths beyond 1m, and the effects of mangrove species, location and environmental context on these stores, is poorly studied. This study investigated these variables at two sites (Gazi and Vanga in the south of Kenya) and used the data to produce a country-specific BGC predictive model for Kenya and map BGC store estimates throughout Kenya at spatial scales relevant for climate change research, forest management and REDD+ (Reduced Emissions from Deforestation and Degradation). The results revealed that mangrove species was the most reliable predictor of BGC; Rhizophora muronata had the highest mean BGC with 1485.5t C ha-1. Applying the species-based predictive model to a base map of species distribution in Kenya for the year 2010 with a 2.5m2 resolution, produced an estimate of 69.41 Mt C (± 9.15 95% C.I.) for BGC in Kenyan mangroves. When applied to a 1992 mangrove distribution map, the BGC estimate was 75.65 Mt C (± 12.21 95% C.I.); an 8.3% loss in BGC stores between 1992 and 2010 in Kenya. The country level mangrove map provides a valuable tool for assessing carbon stocks and visualising the distribution of BGC. Estimates at the 2.5m2 resolution provide sufficient detail for highlighting and prioritising areas for mangrove conservation and restoration.