Developing performance measures of mangrove wetlands using simulation models of hydrology, nutrient biogeochemistry, and community dynamics

The goal of mangrove restoration projects should be to improve community structure and ecosystem function of degraded coastal landscapes. This requires the ability to forecast how mangrove structure and function will respond to prescribed changes in site conditions including hydrology, topography, and geophysical energies. There are global, regional, and local factors that can explain gradients of regulators (e.g., salinity, sulfides), resources (nutrients, light, water), and hydroperiod (frequency, duration of flooding) that collectively account for stressors that result in diverse patterns of mangrove properties across a variety of environmental settings. Simulation models of hydrology, nutrient biogeochemistry, and vegetation dynamics have been developed to forecast patterns in mangroves in the Florida Coastal Everglades. These models provide insight to mangrove response to specific restoration alternatives, testing causal mechanisms of system degradation. We propose that these models can also assist in selecting performance measures for monitoring programs that evaluate project effectiveness. This selection process in turn improves model development and calibration for forecasting mangrove response to restoration alternatives. Hydrologic performance measures include soil regulators, particularly soil salinity, surface topography of mangrove landscape, and hydroperiod, including both the frequency and duration of flooding. Estuarine performance measures should include salinity of the bay, tidal amplitude, and conditions of fresh water discharge (included in the salinity value). The most important performance measures from the mangrove biogeochemistry model should include soil resources (bulk density, total nitrogen, and phosphorus) and soil accretion. Mangrove ecology performance measures should include forest dimension analysis (transects and/or plots), sapling recruitment, leaf area index, and faunal relationships. Estuarine ecology performance measures should include the habitat function of mangroves, which can be evaluated with growth rate of key species, habitat suitability analysis, isotope abundance of indicator species, and bird census. The list of performance measures can be modified according to the model output that is used to define the scientific goals during the restoration planning process that reflect specific goals of the project.

The Role of the Everglades Mangrove Ecotone Region (EMER) in Regulating Nutrient Cycling and Wetland Productivity in South Florida

The authors summarize the main findings of the Florida Coastal Everglades Long-Term Ecological Research (FCE-LTER) program in the EMER, within the context of the Comprehensive Everglades Restoration Plan (CERP), to understand how regional processes, mediated by water flow, control population and ecosystem dynamics across the EMER landscape. Tree canopies with maximum height <3 m cover 49% of the EMER, particularly in the SE region. These scrub/dwarf mangroves are the result of a combination of low soil phosphorus (P < 59 μg P g dw−1) in the calcareous marl substrate and long hydroperiod. Phosphorus limits the EMER and its freshwater watersheds due to the lack of terrigenous sediment input and the phosphorus-limited nature of the freshwater Everglades. Reduced freshwater delivery over the past 50 years, combined with Everglades compartmentalization and a 10 cm rise in coastal sea level, has led to the landward transgression (1.5 km in 54 years) of the mangrove ecotone. Seasonal variation in freshwater input strongly controls the temporal variation of nitrogen and P exports (99%) from the Everglades to Florida Bay. Rapid changes in nutrient availability and vegetation distribution during the last 50 years show that future ecosystem restoration actions and land use decisions can exert a major influence, similar to sea level rise over the short term, on nutrient cycling and wetland productivity in the EMER.

Biogeochemistry of Nitrogen Across the Everglades Landscape

Compared to phosphorus (P), nitrogen (N) has received little attention across the Everglades landscape. Despite this lack of attention, N plays important roles in many Everglades systems, including being a significant pollutant in Florida Bay and the Gulf of Mexico, the limiting nutrient in highly P-impacted areas, and an important substrate for microbial metabolism. Storage and transport of N throughout the Everglades is dominated by organic forms, including peat soils and dissolved organic N in the water column. In general, N sources are highest in the northern areas; however, atmospheric deposition and active N2 fixation by the periphyton components are a significant N source throughout most systems. Many of the processes involved in the wetland N cycle remain unmeasured for most of the Everglades systems. In particular, the lack of in situ rates for N2 fixation and denitrification prevent the construction of system-level budgets, especially for the Southern mangrove systems where N export into Florida Bay is critical. There is also the potential for several novel N processes (e.g., Anammox) with an as yet undetermined importance for nitrogen cycling and function of the Everglades ecosystem. Phosphorus loading alters the N cycle by stimulating organic N mineralization with resulting flux of ammonium and DON, and at elevated P concentrations, by increasing rates of N2 fixation and N assimilation. Restoration of hydrology has a potential for significantly impacting N cycling in the Everglades both in terms of affecting N transport, but also by altering aerobic-anaerobic transitions at the soil-water interface or in areas with seasonal drawdowns (e.g., marl prairies). Based on the authors’ understanding of N processes, much more research is necessary to adequately predict potential impacts from hydrologic restoration, as well as the function of Everglades systems as sinks, sources, and transformers of N in the South Florida landscape.

Ecohydrology, evapotranspiration and hydrogeochemistry of carbonate mangrove wetlands

Coastal environments can be highly susceptible to environmental changes caused by anthropogenic pressures and natural events. Both anthropogenic and natural perturbations may directly affect the amount and the quality of water flowing through the ecosystem, both in the surface and subsurface and can subsequently, alter ecological communities and functions. The Florida Everglades and the Sian Ka'an Biosphere Reserve (Mexico) are two large ecosystems with an extensive coastal mangrove ecotone that represent a historically altered and pristine environment, respectively. Rising sea levels, climate change, increased water demand, and salt water intrusion are growing concerns in these regions and underlies the need for a better understanding of the present conditions. The goal of my research was to better understand various ecohydrological, environmental, and hydrogeochemical interactions and relationships in carbonate mangrove wetlands. A combination of aqueous geochemical analyses and visible and near-infrared reflectance data were employed to explore relationships between surface and subsurface water chemistry and spectral biophysical stress in mangroves. Optical satellite imagery and field collected meteorological data were used to estimate surface energy and evapotranspiration and measure variability associated with hurricanes and restoration efforts. Furthermore, major ionic and nutrient concentrations, and stable isotopes of hydrogen and oxygen were used to distinguish water sources and infer coastal groundwater discharge by applying the data to a combined principal component analysis-end member mixing model. Spectral reflectance measured at the field and satellite scales were successfully used to estimate surface and subsurface water chemistry and model chloride concentrations along the southern Everglades. Satellite imagery indicated that mangrove sites that have less tidal flushing and hydrogeomorphic heterogeneity tend to have more variable evapotranspiration and soil heat flux in response to storms and restoration. Lastly, water chemistry and multivariate analyses indicated two distinct fresh groundwater sources that discharge to the phosphorus-limited estuaries and bays of the Sian Ka'an Biopshere Reserve; and that coastal groundwater discharge was an important source for phosphorus. The results of the study give us a better understanding of the ecohydrological and hydrogeological processes in carbonate mangrove environments that can be then be extrapolated to similar coastal ecosystems in the Caribbean.

Are mangroves in the tropical Atlantic ripe for invasion? Exotic mangrove trees in the forests of South Florida

Abstract Two species of mangrove trees of Indo- Pacific origin have naturalized in tropical Atlantic mangrove forests in South Florida after they were planted and nurtured in botanic gardens. Two Bruguiera gymnorrhiza trees that were planted in the intertidal zone in 1940 have given rise to a population of at least 86 trees growing interspersed with native mangrove species Rhizophora mangle, Avicennia germinans and Laguncularia racemosa along 100 m of shoreline; the population is expanding at a rate of 5.6% year-1. Molecular genetic analyses confirm very low genetic diversity, as expected from a population founded by two individuals. The maximumnumber of alleles at any locus was three, and we measured reduced heterozygosity compared to native-range populations. Lumnitzera racemosa was introduced multiple times during the 1960s and 1970s, it has spread rapidly into a forest composed of native R. mangle, A. germinans, Laguncularia racemosa and Conocarpus erectus and now occupies 60,500 m2 of mangrove forest with stem densities of 24,735 ha-1. We estimate the population growth rate of Lumnitzera racemosa to be between 17 and 23% year-1. Populations of both species of naturalized mangroves are dominated by young individuals. Given the long life and water-dispersed nature of propagules of the two exotic species, it is likely that they have spread beyond our survey area. We argue that the species-depauperate nature of tropical Atlantic mangrove forests and close taxonomic relatives in the more species-rich Indo-Pacific region result in the susceptibility of tropical Atlantic mangrove forests to invasion by Indo-Pacific mangrove species.

Effects of light quantity and quality on early seedling development in the red mangrove Rhizophora mangle L

Seedlings of the red mangrove, Rhizophora mangle L., were grown under light conditions differing in both photosynthetic photon flux density (PPFD) and spectral quality (red:far-red ratio, R:FR). During the first 8 mo of development, parameters of stem, leaf, and root growth were affected by PPFD. Significant responses to lowered R:FR, however, were limited to internode extension. The results are moderately indicative of a strategy to persist in shade, but illustrate the complexity of light responses and suggest that precise categorization as shade-tolerant or -intolerant may be unbefitting for this species at this particular stage of development.

Are mangroves in the tropical Atlantic ripe for invasion? Exotic mangrove trees in the forests of South Florida

Two species of mangrove trees of Indo- Pacific origin have naturalized in tropical Atlantic mangrove forests in South Florida after they were planted and nurtured in botanic gardens. Two Bruguiera gymnorrhiza trees that were planted in the intertidal zone in 1940 have given rise to a population of at least 86 trees growing interspersed with native mangrove species Rhizophora mangle, Avicennia germinans and Laguncularia racemosa along 100 m of shoreline; the population is expanding at a rate of 5.6% year-1. Molecular genetic analyses confirm very low genetic diversity, as expected from a population founded by two individuals. The maximumnumber of alleles at any locus was three, and we measured reduced heterozygosity compared to native-range populations. Lumnitzera racemosa was introduced multiple times during the 1960s and 1970s, it has spread rapidly into a forest composed of native R. mangle, A. germinans, Laguncularia racemosa and Conocarpus erectus and now occupies 60,500 m2 of mangrove forest with stem densities of 24,735 ha-1. We estimate the population growth rate of Lumnitzera racemosa to be between 17 and 23% year-1. Populations of both species of naturalized mangroves are dominated by young individuals. Given the long life and water-dispersed nature of propagules of the two exotic species, it is likely that they have spread beyond our survey area. We argue that the species-depauperate nature of tropical Atlantic mangrove forests and close taxonomic relatives in the more species-rich Indo-Pacific region result in the susceptibility of tropical Atlantic mangrove forests to invasion by Indo-Pacific mangrove species.

The exchange of carbon, nitrogen, and phosphorus in dwarf and fringe mangroves of the oligotrophic southern Everglades

Water management has altered both the natural timing and volume of freshwater delivered to Everglades National Park. This is especially true for Taylor Slough and the C-111 basin, as hypersaline events in Florida Bay have been linked to reduced freshwater flow in this area. In light of recent efforts to restore historical flows to the eastern Everglades, an understanding of the impact of this hydrologic shift is needed in order to predict the trajectory of restoration. I conducted a study to assess the importance of season, water chemistry, and hydrologic conditions on the exchange of nutrients in dwarf and fringe mangrove wetlands along Taylor Slough. I also performed mangrove leaf decomposition studies to determine the contribution of biotic and abiotic processes to mass loss, the effect of salinity and season on degradation rates, and the importance of this litter component as a rapid source of nutrients. ^ Dwarf mangrove wetlands consistently imported total nutrients (C, N, and P) and released NO2− +NO3 −, with enhanced release during the dry season. Ammonium flux shifted from uptake to release over the study period. Dissolved phosphate activity was difficult to discern in either wetland, as concentrations were often below detection limits. Fluxes of dissolved inorganic nitrogen in the fringe wetland were positively related to DIN concentrations. The opposite was found for total nitrogen in the fringe wetland. A dynamic budget revealed a net annual export of TN to Florida Bay that was highest during the wet season. Simulated increases and decreases in freshwater flow yielded reduced exports of TN to Florida Bay as a result of changes in subsystem and water flux characteristics. Finally, abiotic processes yielded substantial nutrient and mass losses from senesced leaves with little influence of salinity. Dwarf mangrove leaf litter appeared to be a considerable source of nutrients to the water column of this highly oligotrophic wetland. To summarize, nutrient dynamics at the subsystem level were sensitive to short-term changes in hydrologic and seasonal conditions. These findings suggest that increased freshwater flow has the potential to lead to long-term, system-level changes that may reach as far as eastern Florida Bay. ^

The successional dynamics of lightning-initiated canopy gaps in the mangrove forests of Shark River, Everglades National Park, USA

Gap succession is a significant determinant of structure and development in most forest communities. Lightning strikes are an important source of canopy gaps in the mangrove forest of Everglades National Park. I investigated the successional dynamics of lightning-initiated canopy gaps to determine their influence on forest stand structure of the mixed mangrove forests ( Rhizophora mangle, Laguncularia racemosa, and Avicennia germinans ) of the Shark River. I measured gap size, gap shape, light environment, soil characteristics, woody debris, and fiddler crab abundance. I additionally measured the vegetative composition in a chronosequences of gap successional stages (new, recruiting, and growing gaps). I recorded survivorship, recruitment, growth and soil elevation dynamics within a subset of new and growing gaps. I determined the relationship between intact forest soil elevation and site hydrology in order to interpret the effects of lightning disturbance on soil elevation dynamics. ^ Gap size averaged 289 ± 20 m2 (± 1SE) and light transmittance decreased exponentially as gaps filled with saplings. Fine woody debris was highest in recruiting gaps. Soil strength was lower in the gaps than in the forest. The abundance of large and medium fiddler crab burrows increased linearly with total seedling abundance. Soil surface elevation declined in newly formed lightning gaps; this loss was due to a combination of superficial erosion (8.5 mm) and subsidence (60.9 mm). A distinct two-cohort recruitment pattern was evident in the seedling/sapling surveys, suggesting a partitioning of the succession between individuals present before and after lightning strike. In new gaps, the seedling recruitment rate was twice as high as in forest and the sapling population increased. At the growing gap stage, R. mangle seedling mortality was 10 times greater and sapling mortality was 13 times greater than recruitment. Growing gaps had reduced seedling stem elongation, sapling growth and adult growth. However, a few individuals (R. mangle saplings) were able to recruit into the adult life stage. In conclusion, the high density of R. mangle seedlings and saplings imply that lightning strike disturbances in these mangrove forests favor their recruitment over that of A. germinans and L. racemosa. ^

Origin, transport and fate of organic matter in Florida Bay: A biomarker record of historical environmental changes

The Everglades are undergoing the world largest wetland restoration project with the aim of returning this system to hydrological conditions in place prior to anthropogenic modifications. Therefore, it is essential to know what these pristine conditions were. In this work, molecular marker (biomarker) distributions and carbon stable isotopic signatures in sediment samples were employed to assess historical environmental changes in Florida Bay over approximately the last 4000 years. Two biomarkers of terrestrial plants, particularly for mangroves (taraxerol and C29 n-alkane), combined with two seagrass proxies (the Paq and the C25/C 27 n-alkan-2-one ratio) revealed a sedimentary environmental shift from freshwater marshes to mangrove swamps and then to seagrass dominated marine ecosystems, likely as a result of sea-level rise in Florida Bay since the Holocene. The maximum values for the Paq and the C 25/C27 n-alkan-2-ones occurred during the 20th century, suggesting that the greatest abundance of seagrass cover is a recent rather than a historical, long-term phenomenon. The greater oscillation in frequency and amplitude for the biomarkers after 1900 potentially reflects an ecosystem under increasing anthropogenic stress. Several algal biomarkers such as C20 highly branched isoprenoids (HBIs), C 25 HBIs and dinosterol indicative of cyanobacteria, diatom and dinoflagellate organic matter inputs respectively, increased dramatically in the latter part of the 20th century and were attributed to recent anthropogenic changes in Florida Bay. ^ The highlight of this work is the development of HBIs as paleo-proxies. As biomarkers of diatoms, the C25 HBIs in the core from the central bay displayed the highest concentration at mid depth, reflecting strong historical inputs of diatom-derived sedimentary OM during that period. In fact, the depth profile of C25 HBIs coincided quite well with historical variations in diatom abundance and variations in diatom species composition in central Florida Bay based on the results of fossil diatom species analysis by microscopy. This study provides evidence that some C25 HBIs can be applied as biomarkers for certain diatom inputs in paleoenvironmental studies. The sources of C20 and C30 HBIs and their potential applicability as paleo-proxies were also investigated and their sources assessed based on their δ13C distributions. ^

Local biomass control on the composition and reactivity of particulate organic matter in aquatic environments

Freshwater ecosystems have been recognized as important components of the global carbon cycle, and the flux of organic matter (OM) from freshwater to marine environments can significantly affect estuarine and coastal productivity. The focus of this study was the assessment of carbon dynamics in two aquatic environments, namely the Florida Everglades and small prairie streams in Kansas, with the aim of characterizing the biogeochemistry of OM. In the Everglades, particulate OM (POM) is mostly found as a layer of flocculent material (floc). While floc is believed to be the main energy source driving trophic dynamics in this oligotrophic wetland, not much is known about its biogeochemistry. The objective of this study was to determine the origin/sources of OM in floc using biomarkers and pigment-based chemotaxonomy to assess specific biomass contributions to this material, on a spatial (freshwater marshes vs. mangrove fringe) and seasonal (wet vs. dry) scales. It was found that floc OM is derived from the local vegetation (mainly algal components and macrophyte litter) and its composition is controlled by seasonal drivers of hydrology and local biomass productivity. Photo-reactivity experiments showed that light exposure on floc resulted in photo-dissolution of POC with the generation of significant amounts of both dissolved OM (DOM) and nutrients (N & P), potentially influencing nutrient dynamics in this ecosystem. The bio-reactivity experiments determined as the amount and rate of CO2 evolution during incubation were found to vary on seasonal and spatial scales and were highly influenced by phosphorus limitation. Not much is known on OM dynamics in small headwater streams. The objective of this study was to determine carbon dynamics in sediments from intermittent prairie streams, characterized by different vegetation cover for their watershed (C4 grasses) vs. riparian zone (C3 plants). In this study sedimentary OM was characterized using a biomarker and compound specific carbon stable isotope approach. It was found that the biomarker composition of these sediments is dominated by higher plant inputs from the riparian zone, although inputs from adjacent prairie grasses were also apparent. Conflicting to some extent with the River Continuum Concept, sediments of the upper reaches contained more degraded OM, while the lower reaches were enriched in fresh material deriving from higher plants and plankton sources as a result of hydrological regimes and particle sorting.

Hurricane disturbance and recovery of energy balance, CO2 fluxes and canopy structure in a mangrove forest of the Florida Everglades

Eddy covariance (EC) estimates of carbon dioxide (CO2) fluxes and energy balance are examined to investigate the functional responses of a mature mangrove forest to a disturbance generated by Hurricane Wilma on October 24, 2005 in the Florida Everglades. At the EC site, high winds from the hurricane caused nearly 100% defoliation in the upper canopy and widespread tree mortality. Soil temperatures down to -50 cm increased, and air temperature lapse rates within the forest canopy switched from statically stable to statically unstable conditions following the disturbance. Unstable conditions allowed more efficient transport of water vapor and CO2 from the surface up to the upper canopy layer. Significant increases in latent heat fluxes (LE) and nighttime net ecosystem exchange (NEE) were also observed and sensible heat fluxes (H) as a proportion of net radiation decreased significantly in response to the disturbance. Many of these impacts persisted through much of the study period through 2009. However, local albedo and MODIS (Moderate Resolution Imaging Spectro-radiometer) data (the Enhanced Vegetation Index) indicated a substantial proportion of active leaf area recovered before the EC measurements began 1 year after the storm. Observed changes in the vertical distribution and the degree of clumping in newly emerged leaves may have affected the energy balance. Direct comparisons of daytime NEE values from before the storm and after our measurements resumed did not show substantial or consistent differences that could be attributed to the disturbance. Regression analyses on seasonal time scales were required to differentiate the storm's impact on monthly average daytime NEE from the changes caused by interannual variability in other environmental drivers. The effects of the storm were apparent on annual time scales, and CO2 uptake remained approximately 250 g C m-2 yr-1 lower in 2009 compared to the average annual values measured in 2004-2005. Dry season CO2 uptake was relatively more affected by the disturbance than wet season values. Complex leaf regeneration dynamics on damaged trees during ecosystem recovery are hypothesized to lead to the variable dry versus wet season impacts on daytime NEE. In contrast, nighttime CO2 release (i.e., nighttime respiration) was consistently and significantly greater, possibly as a result of the enhanced decomposition of litter and coarse woody debris generated by the storm, and this effect was most apparent in the wet seasons compared to the dry seasons. The largest pre- and post-storm differences in NEE coincided roughly with the delayed peak in cumulative mortality of stems in 2007-2008. Across the hurricane-impacted region, cumulative tree mortality rates were also closely correlated with declines in peat surface elevation. Mangrove forest-atmosphere interactions are interpreted with respect to the damage and recovery of stand dynamics and soil accretion processes following the hurricane.

Controls on mangrove forest-atmosphere carbon dioxide exchanges in western Everglades National Park

We report on net ecosystem production (NEP) and key environmental controls on net ecosystem exchange (NEE) of carbon dioxide (CO2) between a mangrove forest and the atmosphere in the coastal Florida Everglades. An eddy covariance system deployed above the canopy was used to determine NEE during January 2004 through August 2005. Maximum daytime NEE ranged from −20 to −25 mmol (CO2) m−2 s−1 between March and May. Respiration (Rd) was highly variable (2.81 ± 2.41 mmol (CO2) m−2 s−1), reaching peak values during the summer wet season. During the winter dry season, forest CO2 assimilation increased with the proportion of diffuse solar irradiance in response to greater radiative transfer in the forest canopy. Surface water salinity and tidal activity were also important controls on NEE. Daily light use efficiency was reduced at high (>34 parts per thousand (ppt)) compared to low (ppt) salinity by 46%. Tidal inundation lowered daytime Rd by ∼0.9 mmol (CO2) m−2 s−1 and nighttime Rd by ∼0.5 mmol (CO2) m−2 s−1. The forest was a sink for atmospheric CO2, with an annual NEP of 1170 ± 127 g C m−2 during 2004. This unusually high NEP was attributed to year‐round productivity and low ecosystem respiration which reached a maximum of only 3 g C m−2 d−1. Tidal export of dissolved inorganic carbon derived from belowground respiration likely lowered the estimates of mangrove forest respiration. These results suggest that carbon balance in mangrove coastal systems will change in response to variable salinity and inundation patterns, possibly resulting from secular sea level rise and climate change. Citation: Barr, J. G., V. Engel, J. D. Fuentes,

Shifting N and P limitation along a north-south gradient of mangrove estuaries in South Florida

A multivariate statistical analysis was applied to a 10 year, multiparameter data set in an effort to describe the spatial dependence and inherent variation of water quality patterns in the mangrove estuaries of Ten Thousand Islands – Whitewater Bay area. Principal component analysis (PCA) of 16 water quality parameters collected monthly resulted in five groupings, which explained 72.5% of the variance of the original variables. The “Organic” component (PCI) was composed of alkaline phosphatase activity, total organic nitrogen, and total organic carbon; the “Dissolved Inorganic N” component (PCII) contained NO 3 − , NO 2 − , and NH 4 + ; the “Phytoplankton” component (PCIII) was made up of total phosphorus, chlorophyll a, and turbidity; dissolved oxygen and temperature were inversely related (PCIV); and salinity and soluble reactive phosphorus made up PCV. A cluster analysis of the mean and SD of PC scores resulted in the spatial aggregation of the 47 fixed stations into six classes having similar water quality, which we defined as: Mangrove Rivers, Whitewater Bay, Gulf Islands, Coot Bay, Blackwater River, and Inland Waterway. Marked differences in physical, chemical, and biological characteristics among classes were illustrated by this technique. Comparison of medians and variability of parameters among classes allowed large scale generalizations as to underlying differences in water quality in these regions. A strong south to north gradient in estuaries from high N - low P to low N - high P was ascribed to marked differences in landuse, freshwater input, geomorphology, and sedimentary geology along this tract. The ecological significance of this gradient discussed along with potential effects of future restoration plans.

Mangrove production and carbon sinks: A revision of global budget estimates

Mangrove forests are highly productive but globally threatened coastal ecosystems, whose role in the carbon budget of the coastal zone has long been debated. Here we provide a comprehensive synthesis of the available data on carbon fluxes in mangrove ecosystems. A reassessment of global mangrove primary production from the literature results in a conservative estimate of ∼218 ± 72 Tg C a−1. When using the best available estimates of various carbon sinks (organic carbon export, sediment burial, and mineralization), it appears that >50% of the carbon fixed by mangrove vegetation is unaccounted for. This unaccounted carbon sink is conservatively estimated at ∼112 ± 85 Tg C a−1, equivalent in magnitude to ∼30–40% of the global riverine organic carbon input to the coastal zone. Our analysis suggests that mineralization is severely underestimated, and that the majority of carbon export from mangroves to adjacent waters occurs as dissolved inorganic carbon (DIC). CO2 efflux from sediments and creek waters and tidal export of DIC appear to be the major sinks. These processes are quantitatively comparable in magnitude to the unaccounted carbon sink in current budgets, but are not yet adequately constrained with the limited published data available so far.