Geology topics

Ilka C. Feller

Publications and source records attributed to Ilka C. Feller.

17 recordsLinked to original sources

USGS2026

Excessive phosphorus loading contributes to future vulnerability of mangrove ecosystems by reducing net ecosystem exchange of carbon

J.N. “Ding” Darling National Wildlife Refuge (DDNWR) is located on Sanibel Island along the southwestern coast of Florida, USA. There, eutrophication attributed to agricultural discharge along the Caloosahatchee River has affected the area’s aquatic habitat. In anticipation of additional nutrient loading, we experimentally fertilized mangrove forests with nitrogen (+N; NH4) and phosphorus (+P; P2O5) for 3 years, and monitored soil and pneumatophore CO2 fluxes and tree sap flow from two mangrove species. Furthermore, we modeled individual tree and stand water use, from which we developed carbon (C) budgets for +N and + P vs. control simulations based on a novel application of water use efficiency conversion. Many of the measured response variables provided hints of subtle changes in response to +P rather than +N, which were enhanced when scaled. From this, we found that additional P loading is expected to reduce both gross and net primary productivity as well as CO2 uptake via net ecosystem exchange of C, likely pressing the system beyond metabolic capacity and leading to a 48–62% decrease in projected lateral C export. Greater eutrophication will likely compound vulnerabilities to sea-level rise submergence, especially where P concentrations are high and already reducing soil surface elevations.

Florida↗

USGS2024

Eutrophication saturates surface elevation change potential in tidal mangrove forests

Coastal mangrove forests are at risk of being submerged due to sea-level rise (SLR). However, mangroves have persisted with changing sea levels due to a variety of biotic and physical feedback mechanisms that allow them to gain and maintain relative soil surface elevation. Therefore, mangrove’s resilience to SLR is dependent upon their ability to build soil elevation at a rate that tracks with SLR, or well-enough to migrate inland. Anthropogenic disturbances, such as altered hydrology and eutrophication, can degrade mangrove forest health and compromise this land building process, placing mangroves at greater risk. Much of Florida’s mangroves are adjacent to highly urbanized areas that produce nutrient-loaded runoff. This study assesses how experimental nutrient inputs in the eutrophic Caloosahatchee Estuary influence the soil surface elevation change (SEC) in two distinct mangrove zones. Annual rates of SEC were reduced by phosphorus additions and differed by mangrove zone, ranging from 0.67 ± 0.59 to 2.13 ± 0.61 and 4.21 ± 0.58 to 6.39 ± 0.59 mm year −1 in the fringe and basin zone, respectively. This suggests that eutrophication can reduce the maximum potential SEC response to SLR and that a mangrove forest’s vulnerability to SLR is not uniform throughout forest but can differ by mangrove zone.

Estuaries and Coasts↗

USGS2023

Temperature thresholds for leaf damage from two extreme freeze events (2018 and 2021) near the northern range limit of black mangroves (Avicennia germinans) in southeastern North America

Extreme winter temperatures govern the northern range limit of black mangroves ( Avicennia germinans ) in southeastern North America. There is a pressing need for studies that advance our understanding of how extreme cold temperature events affect mangroves near their range limits. However, such events are infrequent and challenging to study at regional scales. Here, we compared the damage to mangroves from extreme freeze events in 2018 and 2021, using local data from sites in USA (Florida, Louisiana, and Texas) and northeastern Mexico (Tamaulipas). In 2018, mangrove damage was concentrated in Louisiana and the upper Texas coast, where minimum temperatures ranged from -4 °C to -7 °C. In 2021, damage from a more severe freeze event was concentrated along the central to northern coasts of Texas, where minimum temperatures ranged from -4 °C to -10 °C. We used regional temperature and vegetation data from these events to quantify temperature thresholds for A. germinans leaf damage. Our results indicate that A. germinans leaf damage is likely to occur when temperatures are between -4 °C and -6 °C. These findings help refine temperature thresholds for A. germinans leaf damage and advance understanding of the effects of extreme freeze events on mangrove range expansion. This information is valuable for anticipating future range dynamics in a warming world.

Atlantic Ocean, Gulf of Mexico↗

USGS2023

Rapidly changing range limits in a warming world: Critical data limitations and knowledge gaps for advancing understanding of mangrove range dynamics in the southeastern USA

Climate change is altering species’ range limits and transforming ecosystems. For example, warming temperatures are leading to the range expansion of tropical, cold-sensitive species at the expense of their cold-tolerant counterparts. In some temperate and subtropical coastal wetlands, warming winters are enabling mangrove forest encroachment into salt marsh, which is a major regime shift that has significant ecological and societal ramifications. Here, we synthesized existing data and expert knowledge to assess the distribution of mangroves near rapidly changing range limits in the southeastern USA. We used expert elicitation to identify data limitations and highlight knowledge gaps for advancing understanding of past, current, and future range dynamics. Mangroves near poleward range limits are often shorter, wider, and more shrublike compared to their tropical counterparts that grow as tall forests in freeze-free, resource-rich environments. The northern range limits of mangroves in the southeastern USA are particularly dynamic and climate sensitive due to abundance of suitable coastal wetland habitat and the exposure of mangroves to winter temperature extremes that are much colder than comparable range limits on other continents. Thus, there is need for methodological refinements and improved spatiotemporal data regarding changes in mangrove structure and abundance near northern range limits in the southeastern USA. Advancing understanding of rapidly changing range limits is critical for foundation plant species such as mangroves, as it provides a basis for anticipating and preparing for the cascading effects of climate-induced species redistribution on ecosystems and the human communities that depend on their ecosystem services.

Alabama, Florida, Louisiana, Mississippi, Texas↗

USGS2021

Scientific contributions of the Mangrove Macrobenthos and Management (MMM) conference series, 2000-2019

The conservation of mangrove forests has become an important international policy priority in recent decades, and is mirrored by a large increase in research interest. Multiple disciplines now use mangroves as a study system, from molecular biology to social science. The variety of research conducted in mangroves is exemplified by the Mangrove Macrobenthos & Management (MMM) conference series, the world's largest gathering of researchers and practitioners dedicated to the science and conservation of the mangrove ecosystem. Established in 2000, MMM is a useful barometer with which to identify and measure research trends over the last 20 years. This study describes the history of the MMM conference series, and analyses the research presented in this series as a potential proxy of how the broader mangrove research field has changed through time. Presentations in early MMM conferences were dominated by macrobenthos studies, reflective of the origins of MMM as a forum specifically for mangrove macrobenthos research. However, later conferences have come to reflect the broader interests of the mangrove research field, and have tracked the emergence of blue carbon and other ecosystem services. Mangrove forests continue to be a rich and diverse ecosystem of study, and future MMM conferences will continue to provide a platform for impactful research and management.

Estuarine, Coastal and Shelf Science↗

USGS2019

Temperature thresholds for black mangrove (Avicennia germinans) freeze damage, mortality, and recovery in North America: Refining tipping points for range expansion in a warming climate

Near the tropical‐temperate transition zone, warming winter temperatures are expected to facilitate the poleward range expansion of freeze‐sensitive tropical organisms. In coastal wetlands of eastern and central North America, freeze‐sensitive woody plants (mangroves) are expected to expand northward into regions currently dominated by freeze‐tolerant herbaceous salt marsh plants. To advance understanding of mangrove range expansion, there is a need to refine temperature thresholds for mangrove freeze damage, mortality, and recovery. We integrated data from 38 sites spread across the mangrove range edge in the Gulf of Mexico and Atlantic coasts of North America, including data from a regional collaborative network — the Mangrove Migration Network. In 2018, an extreme freeze event affected 60% of these sites, with minimum temperatures ranging from 0 to ‐7°C. We used temperature and vegetation data from before and after the freeze to quantify temperature thresholds for leaf damage, mortality, and biomass recovery of the black mangrove (Avicennia germinans) — the most freeze‐tolerant mangrove species in North America. For A. germinans individuals near their northern range limit, our results indicate that temperature thresholds for leaf damage are close to ‐4°C, but temperature thresholds for mortality are closer to ‐7°C. Thresholds are expected to be warmer for more southern A. germinans individuals and for the other two common mangrove species in the region (Laguncularia racemosa and Rhizophora mangle). Regenerative buds allowed A. germinans to resprout and recover quickly from aboveground freeze damage. Hence, biomass recovery levels during the first post‐freeze growing season were 90, 78, 62, and 45% for temperatures of ‐4, ‐5, ‐6, and ‐7°C, respectively. Due to a combination of vigorous resprouting and new recruitment from propagules, we expect full recovery at most sites within 1‐3 years, assuming no further freeze events. Synthesis To improve predictions of tropical range expansion in response to climate change, there is a need to better understand tropical species’ responses to winter temperature extremes. Collectively, our results refine temperature thresholds for A. germinans freeze damage, mortality, and recovery, which can improve predictions of mangrove range expansion and coastal wetland ecological transformations in a warming climate.

Florida, Mississippi, Louisiana, Texas↗

USGS2017

The state of the world’s mangroves in the 21st century under climate change

Concerted mangrove research and rehabilitation efforts over the last several decades have prompted a better understanding of the important ecosystem attributes worthy of protection and a better conservation ethic toward mangrove wetlands globally. While mangroves continue to be degraded and lost in specific regions, conservation initiatives, rehabilitation efforts, natural regeneration, and climate range expansion have promoted gains in other areas, ultimately serving to curb the high mangrove habitat loss statistics from the doom and gloom of the 1980s. We highlight those trends in this article and introduce this special issue of Hydrobiologia dedicated to the important and recurring Mangrove and Macrobenthos Meeting. This collection of papers represents studies presented at the fourth such meeting (MMM4) held in St. Augustine, Florida, USA, on July 18–22, 2016. Our intent is to provide a balanced message about the global state of mangrove wetlands by describing recent reductions in net mangrove area losses and highlighting primary research studies presented at MMM4 through a collection of papers. These papers serve not only to highlight on-going global research advancements, but also provide an overview of the vast amount of data on mangrove ecosystem ecology, biology and rehabilitation that emphasizes the uniqueness of the mangrove community.

Hydrobiologia↗

USGS2010

Biocomplexity in mangrove ecosystems

Mangroves are an ecological assemblage of trees and shrubs adapted to grow in intertidal environments along tropical coasts. Despite repeated demonstration of their economic and societal value, more than 50% of the world's mangroves have been destroyed, 35% in the past two decades to aquaculture and coastal development, altered hydrology, sea-level rise, and nutrient overenrichment. Variations in the structure and function of mangrove ecosystems have generally been described solely on the basis of a hierarchical classification of the physical characteristics of the intertidal environment, including climate, geomorphology, topography, and hydrology. Here, we use the concept of emergent properties at multiple levels within a hierarchical framework to review how the interplay between specialized adaptations and extreme trait plasticity that characterizes mangroves and intertidal environments gives rise to the biocomplexity that distinguishes mangrove ecosystems. The traits that allow mangroves to tolerate variable salinity, flooding, and nutrient availability influence ecosystem processes and ultimately the services they provide. We conclude that an integrated research strategy using emergent properties in empirical and theoretical studies provides a holistic approach for understanding and managing mangrove ecosystems.

Annual Review of Marine Science↗

USGS2008

Porewater biogeochemistry and soil metabolism in dwarf red mangrove habitats (Twin Cays, Belize)

Seasonal variability in biogeochemical signatures was used to elucidate the dominant pathways of soil microbial metabolism and elemental cycling in an oligotrophic mangrove system. Three interior dwarf mangrove habitats (Twin Cays, Belize) where surface soils were overlain by microbial mats were sampled during wet and dry periods of the year. Porewater equilibration meters and standard biogeochemical methods provided steady-state porewater profiles of pH, chloride, sulfate, sulfide, ammonium, nitrate/nitrite, phosphate, dissolved organic carbon, nitrogen, and phosphorus, reduced iron and manganese, dissolved inorganic carbon, methane and nitrous oxide. During the wet season, the salinity of overlying pond water and shallow porewaters decreased. Increased rainwater infiltration through soils combined with higher tidal heights appeared to result in increased organic carbon inventories and more reducing soil porewaters. During the dry season, evaporation increased both surface water and porewater salinities, while lower tidal heights resulted in less reduced soil porewaters. Rainfall strongly influenced inventories of dissolved organic carbon and nitrogen, possibly due to more rapid decay of mangrove litter during the wet season. During both times of year, high concentrations of reduced metabolites accumulated at depth, indicating substantial rates of organic matter mineralization coupled primarily to sulfate reduction. Nitrous oxide and methane concentrations were supersaturated indicating considerable rates of nitrification and/or incomplete denitrification and methanogenesis, respectively. More reducing soil conditions during the wet season promoted the production of reduced manganese. Contemporaneous activity of sulfate reduction and methanogenesis was likely fueled by the presence of noncompetitive substrates. The findings indicate that these interior dwarf areas are unique sites of nutrient and energy regeneration and may be critical to the overall persistence and productivity of mangrove-dominated islands in oligotrophic settings. ?? 2008 Springer Science+Business Media B.V.

Biogeochemistry↗

USGS2007

Caribbean mangroves adjust to rising sea level through biotic controls on change in soil elevation

Aim The long-term stability of coastal ecosystems such as mangroves and salt marshes depends upon the maintenance of soil elevations within the intertidal habitat as sea level changes. We examined the rates and processes of peat formation by mangroves of the Caribbean Region to better understand biological controls on habitat stability. Location Mangrove-dominated islands on the Caribbean coasts of Belize, Honduras and Panama were selected as study sites. Methods Biological processes controlling mangrove peat formation were manipulated (in Belize) by the addition of nutrients (nitrogen or phosphorus) to Rhizophora mangle (red mangrove), and the effects on the dynamics of soil elevation were determined over a 3-year period using rod surface elevation tables (RSET) and marker horizons. Peat composition and geological accretion rates were determined at all sites using radiocarbon-dated cores. Results The addition of nutrients to mangroves caused significant changes in rates of mangrove root accumulation, which influenced both the rate and direction of change in elevation. Areas with low root input lost elevation and those with high rates gained elevation. These findings were consistent with peat analyses at multiple Caribbean sites showing that deposits (up to 10 m in depth) were composed primarily of mangrove root matter. Comparison of radiocarbon-dated cores at the study sites with a sea-level curve for the western Atlantic indicated a tight coupling between peat building in Caribbean mangroves and sea-level rise over the Holocene. Main conclusions Mangroves common to the Caribbean region have adjusted to changing sea level mainly through subsurface accumulation of refractory mangrove roots. Without root and other organic inputs, submergence of these tidal forests is inevitable due to peat decomposition, physical compaction and eustatic sea-level rise. These findings have relevance for predicting the effects of sea-level rise and biophysical processes on tropical mangrove ecosystems.

Caribbean Sea↗

USGS2007

Nutrient addition differentially affects ecological processes of Avicennia germinans in nitrogen versus phosphorus limited mangrove ecosystems

Nutrient over-enrichment is a major threat to marine environments, but system-specific attributes of coastal ecosystems may result in differences in their sensitivity and susceptibility to eutrophication. We used fertilization experiments in nitrogen (N)- and phosphorus (P)-limited mangrove forests to test the hypothesis that alleviating different kinds of nutrient limitation may have different effects on ecosystem structure and function in natural systems. We compared a broad range of ecological processes to determine if these systems have different thresholds where shifts might occur in nutrient limitation. Growth responses indicated N limitation in Avicennia germinans (black mangrove) forests in the Indian River Lagoon (IRL), Florida, and P limitation at Twin Cays, Belize. When nutrient deficiency was relieved, A. germinans grew out of its stunted form by increasing wood relative to leaf biomass and shoot length relative to lateral growth. At the P-limited site, P enrichment (+P) increased specific leaf area, N resorption, and P uptake, but had no effect on P resorption. At the N-limited site, +N increased both N and P resorption, but did not alter biomass allocation. Herbivory was greater at the P-limited site and was unaffected by +P, whereas +N led to increased herbivory at the N-limited site. The responses to nutrient enrichment depended on the ecological process and limiting nutrient and suggested that N- versus P-limited mangroves do have different thresholds. +P had a greater effect on more ecological processes at Twin Cays than did +N at the IRL, which indicated that the P-limited site was more sensitive to nutrient loading. Because of this sensitivity, eutrophication is more likely to cause a shift in nutrient limitation at P-limited Twin Cays than N-limited IRL. ?? 2007 Springer Science+Business Media, LLC.

Ecosystems↗

USGS2007

Mangrove recruitment after forest disturbance is facilitated by herbaceous species in the Caribbean

Plant communities along tropical coastlines are often affected by natural and human disturbances, but little is known about factors influencing recovery. We focused on mangrove forests, which are among the most threatened ecosystems globally, to examine how facilitation by herbaceous vegetation might improve forest restoration after disturbance. We specifically investigated whether recovery of mangrove forests in harsh environments is accelerated by nurse plants and whether the beneficial effects are species-specific. Quantification of standardized effects allowed comparisons across performance parameters and over time for: (1) net effect of each herbaceous species on mangrove survival and growth, (2) effects of pre- and post-establishment factors associated with each herbaceous species, and (3) need for artificial planting to enhance growth or survival of mangrove seedlings. Mangrove recruitment in a clear-cut forest in Belize was accelerated by the presence of Sesuvium portulacastrum (succulent forb) and Distichlis spicata (grass), two coastal species common throughout the Caribbean region. The net effect of herbaceous vegetation was positive, but the magnitude of effects on mangrove survival and growth differed by species. Because of differences in their vegetative structure and other features, species effects on mangroves also varied by mechanism: (1) trapping of dispersing propagules (both species), (2) structural support of the seedling (Distichlis), and/or (3) promotion of survival (Sesuvium) or growth (Distichlis) through amelioration of soil conditions (temperature, aeration). Artificial planting had a stronger positive effect on mangrove survival than did edaphic conditions, but planting enhanced mangrove growth more in Sesuvium than in Distichlis patches. Our study indicates that beneficial species might be selected based on features that provide multiple positive effects and that species comparisons may be improved using standardized effects. Our findings are not only relevant to the coastal environments found in the Caribbean region, but our assessment methods may be useful for developing site-specific information to restore disturbed mangrove forests worldwide, especially given the large pool of mangrove associates (>45 genera) available for screening. ?? 2007 by the Ecological Society of America.

Ecological Applications↗

USGS2005

Variation in mangrove forest structure and sediment characteristics in Bocas del Toro, Panama

Mangrove forest structure and sediment characteristics were examined in the extensive mangroves of Bocas del Toro, Republic of Panama. Forest structure was characterized to determine if spatial vegetation patterns were repeated over the Bocas del Toro landscape. Using a series of permanent plots and transects we found that the forests of Bocas del Toro were dominated by Rhizophora mangle with very few individuals of Avicennia germinans and Laguncularia racemosa. Despite this low species diversity, there was large variation in forest structure and in edaphic conditions (salinity, concentration of available phosphorus, Eh and sulphide concentration). Aboveground biomass varied 20-fold, from 6.8 Mg ha-1 in dwarf forests to 194.3 Mg ha-1 in the forests fringing the land. But variation in forest structure was predictable across the intertidal zone. There was a strong tree height gradient from seaward fringe (mean tree height 3.9 m), decreasing in stature in the interior dwarf forests (mean tree height 0.7 m), and increasing in stature in forests adjacent to the terrestrial forest (mean tree height 4.1 m). The predictable variation in forest structure emerges due to the complex interactions among edaphic and plant factors. Identifying predictable patterns in forest structure will aid in scaling up the ecosystem services provided by mangrove forests in coastal landscapes.

Caya Solarte, Isla Colon, Isla Popa, Isla San Cris↗

USGS2004

The effect of nutrient enrichment on growth, photosynthesis and hydraulic conductance of dwarf mangroves in Panamá

1. Dwarf stands of the mangrove Rhizophora mangle L. are extensive in the Caribbean. We fertilized dwarf trees in Almirante Bay, Bocas del Toro Province, north-eastern Panama with nitrogen (N) and phosphorus (P) to determine (1) if growth limitations are due to nutrient deficiency; and (2) what morphological and/or physiological factors underlie nutrient limitations to growth. 2. Shoot growth was 10-fold when fertilized with P and twofold with N fertilization, indicating that stunted growth of these mangroves is partially due to nutrient deficiency. 3. Growth enhancements caused by N or P enrichment could not be attributed to increases in photosynthesis on a leaf area basis, although photosynthetic nutrient-use efficiency was improved. The most dramatic effect was on stem hydraulic conductance, which was increased sixfold by P and 2-5-fold with N enrichment. Fertilization with P enhanced leaf and stem P concentrations and reduced C:N ratio, but did not alter leaf damage by herbivores. 4. Our findings indicate that addition of N and P significantly alter tree growth and internal nutrient dynamics of mangroves at Bocas del Toro, but also that the magnitude, pattern and mechanisms of change will be differentially affected by each nutrient.

Functional Ecology↗

USGS2003

Nitrogen limitation of growth and nutrient dynamics in a disturbed mangrove forest, Indian River Lagoon, Florida

The objectives of this study were to determine effects of nutrient enrichment on plant growth, nutrient dynamics, and photosynthesis in a disturbed mangrove forest in an abandoned mosquito impoundment in Florida. Impounding altered the hydrology and soil chemistry of the site. In 1997, we established a factorial experiment along a tree-height gradient with three zones, i.e., fringe, transition, dwarf, and three fertilizer treatment levels, i.e., nitrogen (N), phosphorus (P), control, in Mosquito Impoundment 23 on the eastern side of Indian River. Transects traversed the forest perpendicular to the shoreline, from a Rhizophora mangle -dominated fringe through an Avicennia germinans stand of intermediate height, and into a scrub or dwarf stand of A. germinans in the hinterland. Growth rates increased significantly in response to N fertilization. Our growth data indicated that this site is N-limited along the tree-height gradient. After 2 years of N addition, dwarf trees resembled vigorously growing saplings. Addition of N also affected internal dynamics of N and P and caused increases in rates of photosynthesis. These findings contrast with results for a R. mangle -dominated forest in Belize where the fringe is N-limited, but the dwarf zone is P-limited and the transition zone is co-limited by N and P. This study demonstrated that patterns of nutrient limitation in mangrove ecosystems are complex, that not all processes respond similarly to the same nutrient, and that similar habitats are not limited by the same nutrient when different mangrove forests are compared.

Florida↗

USGS2003

Nitrogen vs. phosphorus limitation across an ecotonal gradient in a mangrove forest

Mangrove forests are characterized by distinctive tree-height gradients that reflect complex spatial, within-stand differences in environmental factors, including nutrient dynamics, salinity, and tidal inundation, across narrow gradients. To determine patterns of nutrient limitation and the effects of nutrient availability on plant growth and within-stand nutrient dynamics, we used a factorial experiment with three nutrient treatment levels (control, N, P) and three zones along a tree-height gradient (fringe, transition, dwarf) on offshore islands in Belize. Transects were laid out perpendicular to the shoreline across a mangrove forest from a fringe stand along the seaward edge, through a stand of intermediate height, into a dwarf stand in the interior of the island. At three sites, three trees were fertilized per zone for 2 yr. Although there was spatial variability in response, growth by R. mangle was generally nitrogen (N) -limited in the fringe zone; phosphorus (P) -limited in the dwarf zone; and, N- and/or P-limited in the transition zone. Phosphorus-resorption efficiency decreased in all three zones, and N-resorption efficiency increased in the dwarf zone in response to P enrichment. The addition of N had no effect on either P or N resorption efficiencies. Belowground decomposition was increased by P enrichment in all zones, whereas N enrichment had no effect. This study demonstrated that essential nutrients are not uniformly distributed within mangrove ecosystems; that soil fertility can switch from conditions of N to P limitation across narrow ecotonal gradients; and, that not all ecological processes respond similarly to, or are limited by, the same nutrient.

Biogeochemistry↗

USGS2002

Mangrove isotopic (δ15N and δ13C) fractionation across a nitrogen vs. phosphorus limitation gradient

Mangrove islands in Belize are characterized by a unique switching from nitrogen (N) to phosphorus (P) limitation to tree growth from shoreline to interior. Fertilization has previously shown that Rhizophora mangle (red mangrove) fringe trees (5–6 m tall) growing along the shoreline are N limited; dwarf trees ( ≤ 1.5 m tall) in the forest interior are P limited; and transition trees (2–4 m tall) are co-limited by both N and P. Growth patterns paralleled a landward decrease in soil flushing by tides and an increase in bioavailable N, but P availability remained consistently low across the gradient. Stable isotopic composition was measured in R. mangle leaves to aid in explaining this nutrient switching pattern and growth variation. Along control transects, leaf δ15N decreased from +0.10‰ (fringe) to -5.38‰ (dwarf). The δ15N of N-fertilized trees also varied spatially, but the values were consistently more negative (by ~3‰) compared to control trees. Spatial variation in δ15N values disappeared when the trees were fertilized with P, and values averaged +0.12‰, similar to that in control fringe trees. Neither variation in source inputs nor microbial fractionation could fully account for the observed patterns in δ15N . The results instead suggest that the lower δ15N values in transition and dwarf control trees were due to plant fractionation as a consequence of slower growth and lower N demand. P fertilization increased N demand and decreased fractionation. Although leaf δ13C was unaffected by fertilization, values increased from fringe (-28.6‰) to transition (-27.9‰) to dwarf (-26.4‰) zones, indicating spatial variation in environmental stresses affecting stomatal conductance or carboxylation. The results thus suggest an interaction of external supply, internal demand, and plant ability to acquire nutrients under different hydro-edaphic conditions that vary across this tree-height gradient. The findings not only aid in understanding mangrove discrimination of nitrogen and carbon isotopes, but also have implications for identifying nutrient loading and other stress conditions in coastal systems dominated by mangroves.

Twin Cays↗