Environmental Effects of Canopy Gap Formation in High-Rainfall Mangrove Forests1

This study investigated the importance of gap formation in mangrove swamps on the island of Kosrae, Federated States of Micronesia, in order to understand better both natural processes of forest development and the effects of harvesting trees for firewood in these wetlands. Measurements were concentrated in seven plots located near four rivers: three in fringe zones and four in basin zones. Each plot was a cluster of five points and covered an area of ca 1.3 ha. From every point in each of the seven plots, the nearest canopy gap ≥10 m² was located; 25 of the 35 gaps were formed by harvesting. Porewater salinity was significantly higher under the canopy in fringe mangrove forests than in basin mangrove forests. Although gaps were small (mean gap size = 158 m²; median gap size = 92 m²), soil temperatures were significantly higher in gaps of both zones. Soil redox potential was significantly lower and porewater salinity significantly higher in the gaps than under the canopy in the basin zone only. Higher porewater salinity may be attributed to high evaporation rates from the soil and high transpiration rates from trees surrounding gaps. There were significantly more seedlings in gaps than under the canopy only in the fringe zones. Although gap formation alters the soil environment of Kosraean mangrove swamps, high freshwater input may buffer these effects in basin mangrove swamps by reducing porewater salinity. Current harvesting rates do not appear to be changing canopy species composition, but large gaps, especially in mangrove forests in more arid areas, may lead to major changes.     

Soil Respiration and Belowground Carbon Allocation in Mangrove Forests

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

Restoration of Biogeochemical Function in Mangrove Forests

Forest structure of mangrove restoration sites (6 and 14 years old) at two locations (Henderson Creek [HC] and Windstar [WS]) in southwest Florida differed from that of mixed‐basin forests (>50 years old) with which they were once contiguous. However, the younger site (HC) was typical of natural, developing forests, whereas the older site (WS) was less well developed with low structural complexity. More stressful physicochemical conditions resulting from incomplete tidal flushing (elevated salinity) and variable topography (waterlogging) apparently affected plant survival and growth at the WS restoration site. Lower leaf fall and root production rates at the WS restoration site, compared with that at HC were partly attributable to differences in hydroedaphic conditions and structural development. However, leaf and root inputs at each restoration site were not significantly different from that in reference forests within the same physiographic setting. Macrofaunal consumption of tethered leaves also did not differ with site history, but was dramatically higher at HC compared with WS, reflecting local variation in leaf litter processing rates, primarily by snails (Melampus coffeus). Degradation of leaves and roots in mesh bags was slow overall at restoration sites, however, particularly at WS where aerobic decomposition may have been more limited. These findings indicate that local or regional factors such as salinity regime act together with site history to control primary production and turnover rates of organic matter in restoration sites. Species differences in senescent leaf nitrogen content and degradation rates further suggest that restoration sites dominated by Laguncularia racemosa and Rhizophora mangle should exhibit slower recycling of nutrients compared with natural basin forests where Avicennia germinans is more abundant. Structural development and biogeochemical functioning of restored mangrove forests thus depend on a number of factors, but site‐specific as well as regional or local differences in hydrology and concomitant factors such as salinity and soil waterlogging will have a strong influence over the outcome of restoration projects.     

Zonation Patterns of Belizean Offshore Mangrove Forests 41 Years After a Catastrophic Hurricane1

Mangroves are prone to bearing frequently the full brunt of hurricanes and tropical storms. The extent of destruction and early regeneration are widely studied. The purpose of this study was to add a long‐term view of mangrove regeneration and assess the potential effects on mangrove horizontal zonation patterns of catastrophic destruction. Hattie, a category five hurricane, hit the Belizean coast in 1961. It passed directly over the Turneffe Atoll where our study area, Calabash Cay, is located. At four sites on this island, we analyzed mangrove forest structure along transects parallel to the shoreline within zones delineated by species dominance and tree height. We propose an index based on the Simpson index of diversity to express changes in the heterogeneity of the species dominance. Physical–chemical parameters and nutrient availability were also measured. The destruction levels were estimated by analysis of the distribution of diameter at breast heights of the bigger trees in the inland zones. Variations in species dominance among sites and zones could be explained by interactions of various factors. Further, different levels of destruction between the two sides of the island had a significant effect on current patterns of species and structural zonation at Calabash. We conclude that disturbance regime in general should be considered as a factor potentially influencing mangrove horizontal zonation patterns.     

Woody Debris in the Mangrove Forests of South Florida1

Woody debris is abundant in hurricane‐impacted forests. With a major hurricane affecting South Florida mangroves approximately every 20 yr, carbon storage and nutrient retention may be influenced greatly by woody debris dynamics. In addition, woody debris can influence seedling regeneration in mangrove swamps by trapping propagules and enhancing seedling growth potential. Here, we report on line‐intercept woody debris surveys conducted in mangrove wetlands of South Florida 9–10 yr after the passage of Hurricane Andrew. The total volume of woody debris for all sites combined was estimated at 67 m³/ha and varied from 13 to 181 m³/ha depending upon differences in forest height, proximity to the storm, and maximum estimated wind velocities. Large volumes of woody debris were found in the eyewall region of the hurricane, with a volume of 132 m³/ha and a projected woody debris biomass of approximately 36 t/ha. Approximately half of the woody debris biomass averaged across all sites was associated as small twigs and branches (fine woody debris), since coarse woody debris >7.5 cm felled during Hurricane Andrew was fairly well decomposed. Much of the small debris is likely to be associated with post‐hurricane forest dynamics. Hurricanes are responsible for large amounts of damage to mangrove ecosystems, and components of associated downed wood may provide a relative index of disturbance for mangrove forests. Here, we suggest that a fine:coarse woody debris ratio ≤0.5 is suggestive of a recent disturbance in mangrove wetlands, although additional research is needed to corroborate such findings.     

Patterns of local wood use and cutting of Philippine Mangrove Forests

Small-scale wood harvesting from mangrove forests is a commonplace yet barely studied phenomenon. This paper integrates bio-ecological and ethnographic methods to examine local wood use and cutting of mangrove forests in two areas of the Philippines. Findings reveal considerable site variation in cutting intensity, with heavier cutting typically closer to settlements and in forest stands that are not effectively regulated by government or private interests. Overall, cutting is responsible for almost 90% of stem mortality in both natural and plantation forests. Field measurements confirm ethnographic evidence indicating that harvesting for construction wood, but not fuelwood, is both species- and size-selective. Mangrove management and conservation efforts can be made more effective by better understanding how local people are harvesting wood resources from these forests.     

Fatty acids,sterols and hydrocarbons in the leaves from eleven species of mangrove

The fatty acid, sterol and hydrocarbon compositions of the fresh leaves from eleven species of mangroves, cultivated in a shadehouse, are reported. The fatty acid and sterol analyses, whilst generally typical of higher plants, showed several chemotaxonomically significant differences between the species. The epicuticular wax hydrocarbons and fatty acids were low in abundance compared to previous reports of mangrove leaf lipids, which may reflect the importance of environmental influences on this group of compounds. Cluster analysis of selected subsets of the data showed clear chemotaxonomic divisions amongst the mangroves. The results grouped the mangroves into genera, except for the Rhizophora and Ceriops tagal which were not separated, and grouped the family Rhizophoraceae distinct from all other species except Xylocarpus granatum. Avicennia marina var. resinifera was able to be distinguished from Avicennia marina by cluster analysis, supporting its assignment as a distinct variety. The results show that the lipids of mangroves are chemotaxonomically significant.     

Organic Matter Dynamics on the Forest Floor of a Micronesian Mangrove Forest: An Investigation of Species Composition Shifts1

Species composition shifts in mangrove forests may alter organic matter dynamics. The purpose of this study was to predict the effect of species replacements among mangrove trees on organic matter dynamics in a mangrove forest on the island of Kosrae, Federated States of Micronesia. We were particularly interested in elements of the carbon cycle that affect peat accumulation rates, organic matter exports to the estuary and coral reef systems, and soil microbiology. We compared organic matter production and decomposition rates among three mangrove species that commonly grow in similar hydrogeomorphic settings: Rhizophora apiculata BL, which is selectively harvested; Bruguiera gymnorrhiza, which may gradually replace Rhizophora; and Sonneratia alba, which is producing few mature fruits. Sonneratia had significantly higher rates of root production (estimated with ingrowth chambers) than Bruguiera or Rhizophora. Sonneratia foliage had significantly faster decomposition rates and significantly lower lignin:nitrogen ratios than Bruguiera foliage. Live root mass was positively correlated with ingrowth and soil carbon, although soil carbon and ingrowth were not significantly correlated with each other. Humic acid concentrations were significantly higher in Sonneratia rhizospheres than in either Bruguiera or Rhizophora rhizospheres and were positively correlated with root ingrowth. The species changes taking place on Kosrae are likely to result in lower rates of root production and foliage decomposition, and more refractory carbon pools in soil.     

Ecology of Mangrove Growth and Recovery in the Lesser Antilles: State of Knowledge and Basis for Restoration Projects

Whereas the increasing knowledge on tropical coastal wetlands highlights the ecological and economical importance of such ecosystems, anthropogenic activities within the coastal zone have caused substantial, irreversible losses of mangrove areas in the Lesser Antilles during the last decades. Such a paradox gives strength to compensatory policy efforts toward mangrove restoration. We review the available knowledge on the ecology of mangrove growth and recovery in the Lesser Antilles as a contribution to possible restoration projects in such islands. Distribution of species follows a general pattern of seaward/landward zonation according to their respective tolerance to flooding and to pore‐water salinity. An experimental study of seedling growth following simulated oil spill has documented the tolerance of Rhizophora mangle and Avicennia germinans seedlings to oil concentration in soils and the effects of natural biotic and abiotic factors on seedlings growth and survival. Monitoring mangrove recovery following hurricane Hugo has given information on growth patterns, from seedling to sapling stages, according to species and site conditions. Forest recovery was mostly due to pre‐established seedlings. For the large Rhizophora propagules, buoyancy appears to be a quite inefficient way of dispersal far inland from the sea shore or riversides. Causes of recovery failure are discussed. From these results we attempt to answer the questions when, where, how to plant mangroves, and what species to use.     

Role of crabs in the natural regeneration of mangroves in the Sundarbans forest of Bangladesh

Abstract A study in the Bangladesh Sundarbans forest observed the impact of crabs on mangrove regeneration. Crabs were not found to cause any damage to the seedlings of experimental plots. In addition, no morphological damage attributable to crabs was observed in Rhizophora propagules collected from the forest floor. Further studies covering other areas of the forests have been suggested.     

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 sea-level rise.     

Salinity tolerance in some mangrove species from Pakistan

Growth, ionic and water relations of three mangrove species viz. Avicennia marina, Ceriops tagal and Rhizophora mucronata werestudied in different seawater concentrations (0, 25, 50, 75 and 100%).All mangrove species showed optimal growth at 50% seawater. Relativelymore biomass was accumulated by R. mucronata while C. tagalhad the tallest individuals. Tissue water potential became more negativewith the increase in salinity and stomatal conductance was decreased in allplants. Higher stomatal conductance was noted in R. mucronata,followed by A. marina and C. tagal. Sodium and chloride ionsincreased with the increase in salinity and this accumulation was muchhigher in A. marina.     

Disease of Red Mangrove (Rhizophora mangley) in Southwest Puerto Rico Caused by Cytospora rhizophorae1

A high frequency of dieback and mortality was found in Rhizophora mangle (red mangrove) in coastal mangrove forests of southwest Puerto Rico. Eight plots were established in a variety of mangrove environments to assess the extent and severity of the dieback and mortality. The imperfect fungus Cytospora rhizopborae was isolated consistently from stem dieback and associated canker tissues. No other pathogenic microorganisms were isolated. Pathogenicity was demonstrated in greenhouse and field inoculation experiments with C. rhizophorae using young R. mangle seedlings. In field studies of seedlings inoculated, mortality was 33 percent, and greenhouse mortality was as high as 50 percent. Cytospora rhizophorae was isolated consistently from diseased mangrove tissue; the fungus was grown in pure culture and then inoculated into healthy red mangroves, causing cankers. Koch's postulates were fulfilled by infecting R. mangle with C. rhizophorae, producing disease symptoms, and successfully re‐isolating C. rhizophorae from the diseased tissue. This is the first documentation of the pathogenicity of C. rhizophorae associated with mortality in red mangrove. In the coastal mangrove forests of southwest Puerto Rico, C. rhizophorae may play an important role in the frequency of dieback and mortality in R. mangle, producing slow stem diameter growth and causing frequent stem and root wounds.     

Association between pore water sulfide concentrations and the distribution of mangroves

At Humingbird Cay, Exuma, Bahamas, distributions of bothRhizophora mangle (red mangrove) andAvicennia germinans (black mangrove) are closely correlated with amounts of hydrogen sulfide (H₂S) in the soil.R. mangle is primarily distributed within large areas of low to moderate H₂S concentrations (mean= 40 mg/l). H₂S levels underA. germinans are lower (mean = 22 mg/l), but the area immediately beyond their root zone often has extremely high sulfide concentrations (mean= 120 mg/l). These results suggest that past attempts to explain mangrove distribution in terms of monotonic soil gradients, the dispersal characteristics of propagules, and interspecific competition are incomplete, and that it will be necessary to examine the link between soil sulfur chemistry and mangrove distribution more fully.     

The role of freezing in setting the latitudinal limits of mangrove forests

Mangrove trees dominate coastal vegetation in tropical regions, but are completely replaced by herbaceous salt marshes at latitudes above 32 degrees N and 40 degrees S. Because water deficit can increase damage caused by freezing, we hypothesized that mangroves, which experience large deficits as a result of saline substrates, would suffer freeze-induced xylem failure. Vulnerability to freeze-induced xylem embolism was examined in the most poleward mangrove species in North America, in an area where freezing is rare but severe, and in Australia, in an area where freezing is frequent but mild. Percentage loss in hydraulic conductivity was measured following manipulations of xylem tension; xylem sap ion concentration was determined using X-ray microanalysis. Species with wider vessels suffered 60-100% loss of hydraulic conductivity after freezing and thawing under tension, while species with narrower vessels lost as little as 13-40% of conductivity. These results indicate that freeze-induced embolism may play a role in setting the latitudinal limits of distribution in mangroves, either through massive embolism following freezing, or through constraints on water transport as a result of vessel size.     

Leaf Longevity,Construction, and Maintenance Costs of Three Mangrove Species Under Field Conditions

This study assessed the effect of leaf age on construction cost (CC) in the mangrove species Avicennia germinans, Laguncularia racemosa, and Rhizophora mangle growing in their natural habitat. Leaf osmolality values were species-specific, the highest in A. germinans (1 693 mmol kg^–1) and the lowest in L. racemosa (1 270 mmol kg^–1). In the three species, contents of chlorophyll (a+b) (Chl_a+b) and nitrogen (N) per unit of leaf area were maximal in adult leaves and tended to decline with age. Leaf mass to leaf area ratio (LMA) and ash content increased during leaf ageing. Similarly, as leaves aged, a significant increase in leaf construction cost per leaf area (CC_a) was observed, while per leaf mass (CC_m) it remained almost constant, suggesting a sustained production of leaf compounds as leaves became older. CC was positively correlated with LMA and heat of combustion (Hc) per leaf area, suggesting differences among species in the quantity and composition of expensive compounds. Leaf half lifetime (t_0.5) showed contrasting values in the three mangrove species (60, 111, and 160 d in L. racemosa, R. mangle, and A. germinans, respectively). Overall, L. racemosa was the species with less expensive leaves to construct while leaves of A. germinans and R. mangle had the highest CC_m and CC_a, respectively. Leaf longevity was positively correlated with the ratio between CC and maximum photosynthetic rate (P _max), clearly showing the existence of a balance between leaf costs and benefits.     

Two new nitrogen-fixing bacteria from the rhizosphere of mangrove trees: Their isolation, identification and in vitro interaction with rhizosphere Staphylococcus sp.

Abstract Two new diazotrophic bacteria, Listonella anguillarum and Vibrio campbellii , and one non-nitrogen-fixing bacterium, Staphylococcus sp., were isolated from the rhizosphere of mangrove trees. Strains of these newly-defined diazotrophs are known as pathogenic bacteria in fish and shellfish. During the purification of diazotrophic species from the entire rhizosphere population, N₂-fixation of the bacterial mixtures decreased. When grown in vitro in mixed cultures, the non-fixing bacterium Staphylococcus sp. increased the nitrogen-fixing capacity of L. anguillarum by 17% over the pure culture; the nitrogen-fixing capacity per bacterial cell increased 22%. This interaction was not due to a change in O₂ concentration. Staphylococcus sp. decreased the nitrogen-fixing capacity of V. campbellii by 15%.
These findings indicate that (i) other species of rhizosphere bacteria, apart from the common diazotrophic species, should be evaluated for their contribution to the nitrogen-fixation process in mangrove communities; and (ii) the nitrogen-fixing activity detected in the rhizosphere of mangrove plants is probably not the result of individual nitrogen-fixing strains, but the sum of interactions between members of the rhizosphere community.     

The decomposition of mangrove litter in a subtropical mangrove forest

Decomposition of Avicennia marina (Forsk.) Vierh. leaf and woody litter (twigs) was studied using litter bag experiments in a subtropical mangrove forest at two tidal levels (high and low) with different inundation regimes and during two seasons (summer and winter). Losses in dry weight were best described by a single exponential model which showed loss rates of both leaves and twigs were significantly higher low down on the shoreline (greater inundation) and in summer. The time (days) required for the loss of half of the initial dry weight (t ₅₀) was summer: high 59, low 44; winter: high 98, low 78. For twigs the values (days) were summer: high 383, low 179; winter; high 1327, low 1207. There is an exponential relationship between leaf litter t ₅₀'s and latitude which indicates the importance of temperature and therefore season, to the dynamics of organic cycling and export in mangrove systems.