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.

Tree Island Response to Fire and Flooding in the Short-Hydroperiod Marl Prairie Grasslands of the Florida Everglades

Within the marl prairie grasslands of the Florida Everglades, USA, the combined effects of fire and flooding usually lead to very significant changes in tree island structure and composition. Depending on fire severity and post-fire hydroperiod, these effects vary spatially and temporally throughout the landscape, creating a patchy post-fire mosaic of tree islands with different successional states. Through the use of the Normalized Difference Vegetation Index (NDVI) and three predictor variables (marsh water table elevation at the time of fire, post-fire hydroperiod, and tree island size), along with logistic regression analysis, we examined the probability of tree island burning and recovering following the Mustang Corner Fire (May to June 2008) in Everglades National Park. Our data show that hydrologic conditions during and after fire, which are under varying degrees of management control, can lead to tree island contraction or loss. More specifically, the elevation of the marsh water table at the time of the fire appears to be the most important parameter determining the severity of fire in marl prairie tree islands. Furthermore, in the post-fire recovery phase, both tree island size and hydroperiod during the first year after the fire played important roles in determining the probability of tree island recovery, contraction, or loss.

Development and evaluation of a geographic information system based method to estimate flooding susceptibility in an area of Broward County, Florida

The objective of this study was to develop a GIS-based multi-class index overlay model to determine areas susceptible to inland flooding during extreme precipitation events in Broward County, Florida. Data layers used in the method include Airborne Laser Terrain Mapper (ALTM) elevation data, excess precipitation depth determined through performing a Soil Conservation Service (SCS) Curve Number (CN) analysis, and the slope of the terrain. The method includes a calibration procedure that uses "weights and scores" criteria obtained from Hurricane Irene (1999) records, a reported 100-year precipitation event, Doppler radar data and documented flooding locations. Results are displayed in maps of Eastern Broward County depicting types of flooding scenarios for a 100-year, 24-hour storm based on the soil saturation conditions. As expected the results of the multi-class index overlay analysis showed that an increase for the potential of inland flooding could be expected when a higher antecedent moisture condition is experienced. The proposed method proves to have some potential as a predictive tool for flooding susceptibility based on a relatively simple approach.

Vegetation dynamics and their implications for the management of wetlands in the lowlands of Nepal

The maintenance of species richness is often a priority in the management of nature reserves, where consumptive use of resources is generally prohibited. The purpose of this research was to improve management by understanding the vegetation dynamics in the lowlands of Nepal. The objectives were to determine vegetation associations in relation to environments and human-induced disturbances that affect vegetation dynamics on floodplains, where upstream barrages had altered flooding patterns, and consumptive use of plant resources was influencing natural processes. Floodplain vegetation in relation to physical environments and disturbances were studied along transects, perpendicular to the course of the Mahakali River in the western Terai, Nepal. Forest structural changes were studied for three years in ten plots. A randomized split-block experiment with nine burning and grazing treatments was performed in seasonally flooded grasslands. A semi-structured questionnaire was used to assess people's socio-economic status, natural resource use patterns and conservation attitudes. ^ Elevation, soil organic matter, nitrogen, percentage of sand and grazing intensity were significant in delineating herbaceous vegetation assemblages, whereas elevation and livestock grazing were significant in defining forest type boundaries. On the floodplain islands, highly grazed Dalbergia sissoo-Acacia catechu forests were devoid of understory woody vegetation, but the lightly grazed D. sissoo-mixed forests had a well-developed second canopy layer, comprising woody species other than D. sissoo and A. catechu. In grasslands, species richness and biomass production were highest at intermediate disturbance level represented by the lightly grazed and ungrazed early-burned treatments. Ethnicity, education and resource use patterns were important in influencing conservation attitudes. A succession towards the mixed forests would occur in D. sissoo-dominated floodplain forests, where dams and barrages reduce flooding and associated fluvial processes, and if livestock grazing is stopped, as occasionally suggested by nature conservationists. In seasonally flooded grasslands, early burning with moderate grazing would enhance the species diversity and productivity. There is a need to implement a participatory integrated wetland management plan, to include community development, education and off farm income generation, to assure participatory conservation and management of wetlands in Nepal. ^

Short and long hydroperiod Everglades periphyton mats: Community characterization and experimental hydroperiod manipulation

Hydroperiod, or the distribution, duration and timing of flooding affects both plant and animal distributions. The Florida Everglades is currently undergoing restoration that will result in altered hydroperiods. This study was conducted in Everglades National Park to document the variability in periphyton community structure and function between long and short hydroperiod Everglades marshes. Periphyton is an important primary producer and important food resource in the Everglades. Periphyton is also involved in marl soil formation and nutrient cycling. Although periphyton is an important component of the Everglades landscape, little is known about periphyton structural-functional variation between hydroperiods. ^ For this study diatoms, as well as fresh algae slides of diatoms, cyanobacteria and green algae were identified and enumerated. Short verse long hydroperiod soil and water column nutrients were compared. Short and long hydroperiod algal periphyton mat productivity rates were compared using BOD incubations. Experimental manipulations were performed to determine the effects of desiccation duration and rewetting on periphyton productivity, community structure, and nutrient flux. ^ Variation in periphyton community structure was significantly greater between hydroperiods than within hydroperiods. Short and long hydroperiod periphyton mats have the same algal species, it is the distribution and abundance that varies between hydroperiods. Long hydroperiod mats have greater diatom abundance while short hydroperiod mats have greater relative filamentous cyanobacterial abundance. ^ Long hydroperiod mats had greater net primary production (npp) than short hydroperiod mats. Short hydroperiod mats respond to rewetting more rapidly than do long hydroperiod mats. Dry short hydroperiod mats became net primary producers within 24 hours of rehydration. Increasing desiccation duration led to greater cyanobacterial abundance in long hydroperiod mats and decreased diatom abundance in both long and short hydroperiod mats. ^

Rapid responses of vegetation to hydrological changes in Taylor Slough, Everglades National Park, Florida, USA

We analyzed the dynamics of freshwater marsh vegetation of Taylor Slough in eastern Everglades National Park for the 1979 to 2003 period, focusing on cover of individual plant species and on cover and composition of marsh communities in areas potentially influenced by a canal pump station (‘‘S332’’) and its successor station (‘‘S332D’’). Vegetation change analysis incorporated the hydrologic record at these sites for three intervals: pre-S332 (1961–1980), S332 (1980–1999), post-S332 (1999–2002). During S332 and post-S332 intervals, water level in Taylor Slough was affected by operations of S332 and S332D. To relate vegetation change to plot-level hydrological conditions in Taylor Slough, we developed a weighted averaging regression and calibration model (WA) using data from the marl prairies of Everglades National Park and Big Cypress National Preserve. We examined vegetation pattern along five transects. Transects 1–3 were established in 1979 south of the water delivery structures, and were influenced by their operations. Transects 4 and 5 were established in 1997, the latter west of these structures and possibly under their influence. Transect 4 was established in the northern drainage basin of Taylor Slough, beyond the likely zones of influence of S332 and S332D. The composition of all three southern transects changed similarly after 1979. Where muhly grass (Muhlenbergia capillaris var. filipes) was once dominant, sawgrass (Cladium jamaicense), replaced it, while where sawgrass initially predominated, hydric species such as spikerush (Eleocharis cellulosa Torr.) overtook it. Most of the changes in species dominance in Transects 1–3 occurred after 1992, were mostly in place by 1995–1996, and continued through 1999, indicating how rapidly vegetation in seasonal Everglades marshes can respond to hydrological modifications. During the post-S332 period, these long-term trends began reversing. In the two northern transects, total cover and dominance of both muhly grass and sawgrass increased from 1997 to 2003. Thus, during the 1990’s, vegetation composition south of S332 became more like that of long hydroperiod marshes, but afterward it partially returned to its 1979 condition, i.e., a community characteristic of less prolonged flooding. In contrast, the vegetation change along the two northern transects since 1997 showed little relationship to hydrologic status.

Patterns of Root Dynamics in Mangrove Forests Along Environmental Gradients in the Florida Coastal Everglades, USA

Patterns of mangrove vegetation in two distinct basins of Florida Coastal Everglades (FCE), Shark River estuary and Taylor River Slough, represent unique opportunities to test hypotheses that root dynamics respond to gradients of resources, regulators, and hydroperiod. We propose that soil total phosphorus (P) gradients in these two coastal basins of FCE cause specific patterns in belowground biomass allocation and net primary productivity that facilitate nutrient acquisition, but also minimize stress from regulators and hydroperiod in flooded soil conditions. Shark River basin has higher P and tidal hydrology with riverine mangroves, in contrast to scrub mangroves of Taylor basin with more permanent flooding and lower P across the coastal landscape. Belowground biomass (0–90 cm) of mangrove sites in Shark River and Taylor River basins ranged from 2317 to 4673 g m-2, with the highest contribution (62–85%) of roots in the shallow root zone (0–45 cm) compared to the deeper root zone (45–90 cm). Total root productivity did not vary significantly among sites and ranged from 407 to 643 g m-2 y-1. Root production in the shallow root zone accounted for 57–78% of total production. Root turnover rates ranged from 0.04 to 0.60 y-1 and consistently decreased as the root size class distribution increased from fine to coarse roots, indicating differences in root longevity. Fine root biomass was negatively correlated with soil P density and frequency of inundation, whereas fine root turnover decreased with increasing soil N:P ratios. Lower P availability in Taylor River basin relative to Shark River basin, along with higher regulator and hydroperiod stress, confirms our hypothesis that interactions of stress from resource limitation and long duration of hydroperiod account for higher fine root biomass along with lower fine root production and turnover. Because fine root production and organic matter accumulation are the primary processes controlling soil formation and accretion in scrub mangrove forests, root dynamics in the P-limited carbonate ecosystem of south Florida have a major controlling role as to how mangroves respond to future impacts of sealevel rise.

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.

Responses of neotropical mangrove seedlings grown in monoculture and mixed culture under treatments of hydroperiod and salinity

We investigated the combined effects of salinity and hydroperiod on seedlings of Rhizophora mangle and Laguncularia racemosa grown under experimental conditions of monoculture and mixed culture by using a simulated tidal system. The objective was to test hypotheses relative to species interactions to either tidal or permanent flooding at salinities of 10 or 40 g/l. Four-month-old seedlings were experimentally manipulated under these environmental conditions in two types of species interactions: (1) seedlings of the same species were grown separately in containers from September 2000 to August 2001 to evaluate intraspecific response and (2) seedlings of each species were mixed in containers to evaluate interspecific, competitive responses from August 2002 to April 2003. Overall, L. racemosa was strongly sensitive to treatment combinations while R. mangle showed little effect. Most plant responses of L. racemosa were affected by both salinity and hydroperiod, with hydroperiod inducing more effects than salinity. Compared to R. mangle, L. racemosa in all treatment combinations had higher relative growth rate, leaf area ratio, specific leaf area, stem elongation, total length of branches, net primary production, and stem height. Rhizophora mangle had higher biomass allocation to roots. Species growth differentiation was more pronounced at low salinity, with few species differences at high salinity under permanent flooding. These results suggest that under low to mild stress by hydroperiod and salinity, L. racemosa exhibits responses that favor its competitive dominance over R. mangle. This advantage, however, is strongly reduced as stress from salinity and hydroperiod increase.

Fish population dynamics in a seasonally varying wetland

Small fishes in seasonally flooded environments such as the Everglades are capable of spreading into newly flooded areas and building up substantial biomass. Passive drift cannot account for the rapidity of observed population expansions. To test the ‘reaction–diffusion’ mechanism for spread of the fish, we estimated their diffusion coefficient and applied a reaction–diffusion model. This mechanism was also too weak to account for the spatial dynamics. Two other hypotheses were tested through modeling. The first—the ‘refuge mechanism’—hypothesizes that small remnant populations of small fishes survive the dry season in small permanent bodies of water (refugia), sites where the water level is otherwise below the surface. The second mechanism, which we call the ‘dynamic ideal free distribution mechanism’ is that consumption by the fish creates a prey density gradient and that fish taxis along this gradient can lead to rapid population expansion in space. We examined the two alternatives and concluded that although refugia may play an important role in recolonization by the fish population during reflooding, only the second, taxis in the direction of the flooding front, seems capable of matching empirical observations. This study has important implications for management of wetlands, as fish biomass is an essential support of higher trophic levels.

Hydrologic Dynamics of the Ground-Water-Dependent Sian Ka’an Wetlands, Mexico, Derived from InSAR and SAR Data

The 5,280 km2 Sian Ka’an Biosphere Reserve includes pristine wetlands fed by ground water from the karst aquifer of the Yucatan Peninsula, Mexico. The inflow through underground karst structures is hard to observe making it difficult to understand, quantify, and predict the wetland dynamics. Remotely sensed Synthetic Aperture Radar (SAR) amplitude and phase observations offer new opportunities to obtain information on hydrologic dynamics useful for wetland management. Backscatter amplitude of SAR data can be used to map flooding extent. Interferometric processing of the backscattered SAR phase data (InSAR) produces temporal phase-changes that can be related to relative water level changes in vegetated wetlands. We used 56 RADARSAT-1 SAR acquisitions to calculate 38 interferograms and 13 flooding maps with 24 day and 48 day time intervals covering July 2006 to March 2008. Flooding extent varied between 1,067 km2 and 2,588 km2 during the study period, and main water input was seen to take place in sloughs during October–December. We propose that main water input areas are associated with water-filled faults that transport ground water from the catchment to the wetlands. InSAR and Landsat data revealed local-scale water divides and surface water flow directions within the wetlands.

Patterns of nutrient exchange in a riverine mangrove forest in the Shark River Estuary, Florida, USA

This study aimed to evaluate tidal and seasonal variations in concentrations and fluxes of nitrogen (NH4 +, NO2+NO3, total nitrogen) and phosphorus (soluble reactive phosphorus, total phosphorus) in a riverine mangrove forest using the flume technique during the dry (May, December 2003) and rainy (October 2003) seasons in the Shark River Estuary, Florida. Tidal water temperatures during the sampling period were on average 29.4 (± 0.4) oC in May and October declining to 20 oC (± 4) in December. Salinity values remained constant in May (28 ± 0.12 PSU), whereas salinity in October and December ranged from 6‒21 PSU and 9‒25 PSU, respectively. Nitrate + nitrite (N+N) and NH4+ concentrations ranged from 0.0 to 3.5 μM and from 0 to 4.8 μM throughout the study period, respectively. Mean TN concentrations in October and December were 39 (±0.8) μM and 37 (±1.5) μM, respectively. SRP and N+N concentrations in the flume increased with higher frequency in flooding tides. TP concentrations ranged between 0.2‒2.9 μM with higher concentrations in the dry season than in the rainy season. Mean concentrations were <1. 5 μM during the sampling period in October (0.75 ± 0.02) and December (0.76 ± 0.01), and were relatively constant in both upstream and downstream locations of the flume. Water residence time in the flume (25 m2) was relatively short for any nutrient exchange to occur between the water column and the forest floor. However, the distinct seasonality in nutrient concentrations in the flume and adjacent tidal creek indicate that the Gulf of Mexico is the main source of SRP and N+N into the mangrove forest.

Modeling seasonal dynamics of small fish cohorts in fluctuating freshwater marsh landscapes

Small-bodied fishes constitute an important assemblage in many wetlands. In wetlands that dry periodically except for small permanent waterbodies, these fishes are quick to respond to change and can undergo large fluctuations in numbers and biomasses. An important aspect of landscapes that are mixtures of marsh and permanent waterbodies is that high rates of biomass production occur in the marshes during flooding phases, while the permanent waterbodies serve as refuges for many biotic components during the dry phases. The temporal and spatial dynamics of the small fishes are ecologically important, as these fishes provide a crucial food base for higher trophic levels, such as wading birds. We develop a simple model that is analytically tractable, describing the main processes of the spatio-temporal dynamics of a population of small-bodied fish in a seasonal wetland environment, consisting of marsh and permanent waterbodies. The population expands into newly flooded areas during the wet season and contracts during declining water levels in the dry season. If the marsh dries completely during these times (a drydown), the fish need refuge in permanent waterbodies. At least three new and general conclusions arise from the model: (1) there is an optimal rate at which fish should expand into a newly flooding area to maximize population production; (2) there is also a fluctuation amplitude of water level that maximizes fish production, and (3) there is an upper limit on the number of fish that can reach a permanent waterbody during a drydown, no matter how large the marsh surface area is that drains into the waterbody. Because water levels can be manipulated in many wetlands, it is useful to have an understanding of the role of these fluctuations.

Leaf gas exchange characteristics of three neotropical mangrove species in response to varying hydroperiod

We determined how different hydroperiods affected leaf gas exchange characteristics of greenhouse-grown seedlings (2002) and saplings (2003) of the mangrove species Avicennia germinans (L.) Stearn., Laguncularia racemosa (L.) Gaertn. f., and Rhizophora mangle L. Hydroperiod treatments included no flooding (unflooded), intermittent flooding (intermittent), and permanent flooding (flooded). Plants in the intermittent treatment were measured under both flooded and drained states and compared separately. In the greenhouse study, plants of all species maintained different leaf areas in the contrasting hydroperiods during both years. Assimilation-light response curves indicated that the different hydroperiods had little effect on leaf gas exchange characteristics in either seedlings or saplings. However, short-term intermittent flooding for between 6 and 22 days caused a 20% reduction in maximum leaf-level carbon assimilation rate, a 51% lower light requirement to attain 50% of maximum assimilation, and a 38% higher demand from dark respiration. Although interspecific differences were evident for nearly all measured parameters in both years, there was little consistency in ranking of the interspecific responses. Species by hydroperiod interactions were significant only for sapling leaf area. In a field study, R. mangle saplings along the Shark River in the Everglades National Park either demonstrated no significant effect or slight enhancement of carbon assimilation and water-use efficiency while flooded. We obtained little evidence that contrasting hydroperiods affect leaf gas exchange characteristics of mangrove seedlings or saplings over long time intervals; however, intermittent flooding may cause short-term depressions in leaf gas exchange. The resilience of mangrove systems to flooding, as demonstrated in the permanently flooded treatments, will likely promote photosynthetic and morphological adjustment to slight hydroperiod shifts in many settings.

Evaluating the relative contributions of hydroperiod and soil fertility on growth of south Florida mangroves

Low and high water periods create contrasting challenges for trees inhabiting periodically flooded wetlands. Low to moderate flood durations and frequencies may bring nutrient subsidies, while greater hydroperiods can be energetically stressful because of oxygen deficiency. We tested the hypothesis that hydroperiod affects the growth of mangrove seedlings and saplings in a greenhouse experiment by varying flood duration while keeping salinity and soil fertility constant. We measured the growth of mangrove trees along a hydroperiod gradient over a two-year period by tracking fine-scale diameter increment. Greenhouse growth studies indicated that under a full range of annual flood durations (0–8760 h/year), hydroperiod alone exerted a significant influence on growth for one species, Laguncularia racemosa, when flooding was imposed for two growing seasons. Field evaluations, on the other hand, indicated that increased flood duration may provide nutrient subsidies for tree growth. Diameter growth was related curvilinearly to site hydroperiod, including flood duration and frequency, as well as to salinity and soil fertility. An analysis of soil physico-chemical parameters suggests that phosphorus fertility, which was also linked directly to hydroperiod, is likely to influence growth on south Florida mangrove sites. The physical removal of phosphorus by greater flood frequencies from upland sources and/or addition of phosphorus from tidal flooding balanced against increased soil aeration and reduced water deficits may be an extremely important growth determinant for south Florida mangroves.